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VOL. 101 



JANUARY 1999 



NO. 1 

(ISSN 0013-8797) 



PROCEEDINGS' 

of the \ 




ENTOMOLOGICAL SOCI 



of WASHINGTON 

PUBLISHED 
QUARTERLY 




CONTENTS 

ADAMSKI, DAVID — Blastobasis graminea, new species (Lepidoptera: Gelechioidea: Coleo- 

phoridae: Blastobasinae), a stem borer of sugar cane in Colombia and Venezuela 164 

BAIER, TEVIS M. — A revision of the flavidus group of the genus Chrysops Meigen (Diptera: 

Tabanidae) 175 

BATRA, SUZANNE W. T. — Biology of Andrena (Scrapteropsis) fenningeri Viereck (Hyme- 

noptera: Andrenidae), harbinger of spring 1 06 

BRAILOVSKY, HARRY and GERASIMOS CASSIS— New genus and new species of Amorbini 

(Heteroptera: Coreidae) from Australia 69 

BREIDENBAUGH. MARK S. and BRADLEY A. MULLENS— Two new western Nearctic 

Culicoides Latreille (Diptera: Ceratopogonidae) descibed from all stages 149 

CARVALHO, CLAUDIO JOSE BARROS DE— Revision, cladistics and biogeography of the 

Neotropical genus Souzalopesmyia Albuquerque (Diptera: Muscidae) 1 23 

EASTON, EMMETT R. and WING-WAH PUN— Observations on twelve families of Homoptera 

in Macau, southeastern China, from 1 989 to the present 99 

GAGNE, RAYMOND J. and JOHN F. DOANE— The larval instars of the wheat midge, Sito- 

diplosis mosellana (Gehin) (Diptera: Cecidomyiidae) 57 

GOEDEN, RICHARD D. and JEFFREY A. TEERINK— Life history and description of imma- 
ture stages of Trupanea arizonensis Malloch (Diptera: Tephritidae) on Trixis californica 
Kellogg var. californica (Asteraceae) in southern California 75 

HARRIS, S. C. and L. J. DAVENPORT— New species of Hydroptilidae (Trichoptera) from the 

Amazon region of northeastern Peru 26 

JENSEN, ANDREW S. and MANYA B. STOETZEL— An examination of the North American 
aphid species currently placed in Ovatus van der Goot (Hemiptera: Aphididae) with the 
description of a new genus 39 

JOHNSON, NORMAN F., LUCIANA MUSETTI. JAMES B. JOHNSON, and KERRY KA- 

TOVICH — The larva of Pelecinus polyturator (Drury) (Hymenoptera: Pelecinidae) 64 

{Continued on back cover) 



THE 

ENTOMOLOGICAL SOCIETY 

OF WASHINGTON 

Organized March 12, 1884 

OFFICERS FOR 1999 

Michael E. Schauff, President Michael G. Pogue, Treasurer 

David G. Furth, President-Elect John W. Brown, Program Chair 

Stuart H. McKamey, Recording Secretary Steven W. Lingafelter, Membership Chair 

Mollis B. Williams, Corresponding Secretary Warren E. Steiner, Past President 
Andrew S. Jensen, Custodian 

David R. Smith, Editor 

Publications Committee 
Thomas J. Henry Wayne N. Mathis 

Gary L. Miller, Book Review Editor 



Honorary President 
Louise M. Russell 



Honorary Members 
Alan Stone Karl V. Krombein Ronald W. Hodges 



All correspondence concerning Society business should be mailed to the appropriate officer at the following 
address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- 
ington, D.C. 20560-0168. 

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tion, on the first Thursday of each month from October to June, inclusive, at 7:30 P.M. Minutes of meetings 
are published regularly in the Proceedings. 

MEMBERSHIP. — Members shall be persons who have demonstrated interest in the science of entomology. 
Annual dues for members are $25.00 (U.S. currency). 

PROCEEDINGS.— The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- 
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The Society does not exchange its publications for those of other societies. 

PLEASE SEE PP. 599-600 OF THE JULY 1998 ISSUE FOR INFORMATION REGARDING 
PREPARATION OF MANUSCRIPTS. 

STATEMENT OF OWNERSHIP 

Title of Publication: Proceedings of the Entomological Society of Washington. 

Frequency of Issue: Quarterly (January, April, July, October). 

Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of 
Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- 
ington, D.C. 20560-0168. 

Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, 
Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. 

Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- 
West, Beltsville, MD 20705. 

Managing Editor and Known Bondholders or other Security Holders: none. 

This issue was mailed 19 January 1999 

Second Class Postage Paid at Washington, DC. and additional mailing office. 
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044. USA 

@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999. pp. 1-25 

EGG ARCHITECTURE OF NAUCORIDAE (HETEROPTERA): INTERNAL 
AND EXTERNAL STRUCTURE OF THE CHORION AND MICROPYLE 

Robert W. Sites and Becky J. Nichols 

Enns Entomology Museum, Department of Entomology, University of Missouri, Co- 
lumbia, MO 65211, U.S.A. (e-mail: bugs@showme.missouri.edu) 



Abstract. — The chorion and micropyles of 27 species of Naucoridae, including 21 spe- 
cies of Ambrysus, are described and each description is supported with scanning electron 
micrographs. Photomicrographs of thick sections through the chorion or micropyle of 10 
of these species also are presented. Chorionic sculpturing differs interspecifically in Am- 
brysus and based on the species studied, eggs of Ambrysus, Limnocoris, and Pelocoris 
have 2-3 micropyles. 

Key Words: Egg, chorion, micropyle, Naucoridae 



The Naucoridae (sensu lato), or creeping 
water bugs, comprise 394 described species 
worldwide (see La Rivers 1971, 1974, 
1976; Polhemus and Polhemus 1982, 1988, 
1994; Nieser et al. 1993; Liu and Zheng 
1994; Polhemus 1994; Nieser and Chen 
1996). As is typical among insect families, 
the adults have received the greatest 
amount of morphological research; accord- 
ingly, taxonomic treatments emphasize 
adult characters. Although nymphal stages 
have been described for several species of 
Naucoridae, details of egg structure largely 
have been ignored. Line diagrams from 
light microscopy have been presented for a 
few species; however, elucidation of fine 
detail is not possible with this technique. 
Recent reports for several species [Ambry- 
sus lunatus Usinger (Sites and Nichols 
1990), Cryphocricos hungerfordi Usinger 
(Sites and Nichols 1993), Pelocoris poeyi 
Guerin Meneville (Sites 1991), and several 
species of South American Ambrysus and 
Pelocoris (Lopez Ruf 1989)] have included 
scanning electron micrographs that have re- 
vealed interspecific differences in chorionic 
patterns. 



Eggs of most naucorid species, for which 
oviposition is known, either are adhered to 
plants (exophytic oviposition) or to rock 
substrata (Hinton 1981). For example, eggs 
of A. lunatus are adhered to plants (Sites 
and Nichols 1990), Ambrysus mormon 
Montandon to pebbles (Usinger 1946), 
Aphelocheirus aestivalis (Fabricius) proba- 
bly to rocks (Larsen 1927), C. hungerfordi 
to rocks (Sites and Nichols 1993), Lacco- 
coris limigenus Stal to hard substrata 
(Clarke and Baroudy 1990), Naucoris ma- 
culatus F. to plants (Lebrun 1960), and Pel- 
ocoris femoratus (Palisot de Beauvois) to 
plants (Torre Bueno 1903, Hungerford 
1927, McPherson et al. 1987). More spe- 
cifically, P. femoratus eggs are glued to leaf- 
lets of Nitella and other aquatic plants with 
a "fairly generous quantity of white adhe- 
sive" (Hungerford 1927). In contrast, the 
oviposition of Ilyocoris cimicoides (L.) is 
endophytic, inserting eggs into submergent 
plant tissue (Cobben 1968), such as into 
stems of Ranunculus or water peppermint 
(Rawat 1939). 

Eclosion occurs through the anterior pole 
and, generally, a crescentic slit is made 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



through the chorion. In Coleopterocoris 
kleerekopeh Hungerford and species of 
Cryphocricos, a predetermined fracture Une 
exists, and in /. cimicoides, a well-defined 
operculum faces the water, whereas the re- 
mainder of the egg is embedded within the 
plant (see Rawat 1939). 

Internally, the chorion of Ilyocoris and 
Cryphocricos is bilayered (Cobben 1968 
and Sites and Nichols 1993, respectively), 
with a thick chorionic outer layer, which is 
perforated by pore canals, and a thinner, un- 
perforated chorionic inner layer. Externally, 
scanning electron micrographs of six spe- 
cies of Pelocoris and two species of Am- 
brysus from Argentina revealed that the 
chorionic surface differed among species 
(Lopez Ruf 1989). For the two species of 
Ambrysus, interspecific internal differentia- 
tion was subtle; however, marked differenc- 
es existed externally. Therefore, Lopez Ruf 
(1989) suggested that the external chorionic 
pattern is valuable as a taxonomic character 
and that internal chorionic attributes may be 
useful at the generic level. 

Presented herein are scanning electron 
micrographs of the chorion and micropyles 
of 21 species oi Ambrysus (subfamily Cry- 
phocricinae) and selected species represent- 
ing six additional genera and five additional 
subfamilies. Photomicrographs of thick sec- 
tions of the micropyle and chorion for some 
species also are presented. Egg morphology 
is described for each species. 

Materials and Methods 

Eggs were obtained by both oviposition 
and dissection. Oviposited eggs were pre- 
ferred for examination because they were 
fully developed structurally. Thus, we 
brought live female naucorids into the lab- 
oratory and maintained them individually in 
glass petri dishes with enough water to sub- 
merge them. To provide a potential ovipo- 
sition substrate, an aquatic plant stem (usu- 
ally Justicia americana) was placed in each 
petri dish and live food provided for each 
naucorid generally as one corixid {Corisel- 
la, Ramphocorixa, Sigara) per naucorid. 



daily. Most naucorids, including members 
of those species inhabiting lotic environ- 
ments, oviposited on plants or on the dish. 
For the several species that did not oviposit 
in the laboratory or for which we did not 
have live specimens, eggs were dissected 
from females preserved in alcohol. Eggs 
were taken from only the common oviduct 
or vagina (sometimes erroneously referred 
to as ovarian eggs), rather than from the 
ovarioles, and were considered to be struc- 
turally well-developed because they were 
near the end of the reproductive tract. To 
allow a rapid evaluation of the reproductive 
tract to determine position of eggs and to 
minimize external damage to the specimen, 
a dissection technique was developed. With 
the insect in alcohol and ventral side up, the 
tip of a pair of jeweler's forceps was in- 
serted into the membrane of the lateral mar- 
gin of the 7th abdominal segment. By mov- 
ing the forceps anteriorly, the sternum and 
laterotergites were separated from the terga 
of segments 4-7. The venter then was 
pulled laterally, separating along segmental 
sutures, thereby exposing the abdominal 
cavity. Eggs were gently removed from the 
abdomen with forceps and kept in 3.7 ml 
snap-cap glass vials in 80% ethyl alcohol. 

Dissected eggs often had tissue from the 
reproductive tract adhered to them. To re- 
move this tissue, an ultrasonic cleaner was 
used, which had a peak output of 40 watts 
and a frequency of 60 Hz. Vials containing 
eggs in ethyl alcohol were placed in the 
cleaner with a small amount of water for 
ca. 10 minutes: the exact amount of time 
depended on the amount of tissue to be re- 
moved. Various solublizers [e.g., Triton X- 
100, sodium dodecyl sulfate (SDS)] were 
ineffective in removing tissue. Subsequent- 
ly, the eggs were examined under a micro- 
scope and any remaining tissue was care- 
fully removed. Eggs then were transferred 
to fresh 80% ethyl alcohol. 

To prepare eggs for critical point drying, 
both dissected and oviposited eggs in 80% 
ethyl alcohol were fixed with 2% glutaral- 
dehyde in 0.1 M phosphate buffer, washed 



VOLUME 101. NUMBER 1 





ll@9 



Fig. 1. Ambrysiis circiiincinctiis. cross section 
through micropylar plug distad of base. 



in 0.1 M phosphate buffer, and dehydrated 
through a graded alcohol series to 100% 
ethyl alcohol, and then placed into 100% 
acetone. Eggs were then critical point dried 
and sputter-coated. For light microscopy, 
the above procedure was followed through 
the glutaraldehyde primary fixation. After 
washing in 0.1 M phosphate buffer, eggs 
were subjected to a secondary fixation in 
2% osmium tetroxide in 0.1 M phosphate 
buffer. Subsequent dehydration techniques 
were as for the scanning microscopy prep- 
aration. Eggs were embedded in Spurr's 
resin, sectioned with a diamond knife, 
slide-mounted, and stained with toluidine 
blue. Voucher specimens of adults and eggs 
are housed in the Enns Entomology Muse- 
um, University of Missouri-Columbia. 

Descriptions of Eggs 

Little detail of the micropylar plug of 
most genera is evident externally that 
would allow the determination of the num- 
ber of micropyles contained therein. Cross 
sections taken at levels above the base of 
the plug reveal numerous canals, including 
transverse canals, which represent convo- 
lutions of the same canal(s) throughout the 
plug (Fig. 1). However, the number of mi- 
cropyles can be observed by sectioning 
through the base of the plug where it enters 
the egg. Thus, photomicrographs of thick 



sections taken through the base of the plug 
are presented for some species. 

Citations for the original description and 
subsequent descriptions of adults or imma- 
ture stages are given for each species. Also 
given are collecting localities of females 
from which eggs were obtained, egg mea- 
surements (mm ± SE), sample size, and 
method of obtainment (i.e., oviposited or 
dissected). For all naucorid species exam- 
ined, egg color was creamy white to beige. 

Subfamily Cryphocricinae Montandon 

1897a 

Genus Ambrysus Stal 1862 

Ambrysus (Acyttarus) fimebris La Rivers 

(Figs. 2-5) 

Ambrysus funebris La Rivers 1948a: 103- 

107. 
USA: California, Death Valley National 

Monument 
Length, 1.04; width, 0.38; n = 1; dissected. 

Overall appearance elongate with round- 
ed, asymmetrical ends. Reticulation pattern 
consisting generally of pentagonal to hep- 
tagonal units, delimited by distinctly raised 
lines (Fig. 2). Within each unit, poorly de- 
fined, low, irregular protruberances. Aero- 
pyles small, distinct, numerous, more than 
50 per unit (Fig. 3). Anterior pole with re- 
ticulation and aeropyles lacking, with amor- 
phous micropylar plug set in shallow con- 
cavity (Fig. 4). 

Exochorion thicker than endochorion. 
Pore canals widest at base (Fig. 5). 



Ambrysus (SyncoUus) circumcinctus 
Montandon 
(Figs. 6-9) 

Ambrysus circumcinctus Montandon 1910: 

442-444. 
USA: Texas, Kimble Co., Junction 
Length, 1.11 ± 0.01; width, 0.48 ± 0.01; n 

= 10; dissected. 

Overall appearance elongate-oval. Retic- 
ulation pattern only faintly visible as im- 
pressed lines. Within each unit formed by 




PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

■ -• ■-'•-;^ • ;■'-■ ■':.•■* i'-^':n:-."^"-^^-^ 
I , ^ ; ' • ■■ • ' ••^« - 1 ^■\- • ' -\ ' ^ ' j;. V * 

■ '.';.'*''. .''■-■ *. '..".'*/,' '.V'"^'* f. T --- v 

I ^'C'^-' ^ ;■ ,^ • ^;>.- ■•■■ w ;--;♦-. '.-^;\ •*-.*,'.;% 






1 




Figs. 2-7. 2-5, Ambrysus funebris. 2, 3, Chorion surface. 4, Micropylar plug. 5, Chorion section. 6. 7, /I. 
■tus. 6, Chorion surface. 7, Micropylar plug. 



circumcmci 



pattern, 5-8 aeropyles set in deep individual Section through base of micropylar plug 

sockets (Fig. 6). Anterior pole with reticu- with two micropyles (Fig. 8). Protuberances 

lation pronounced with raised lines and other than micropyle lacking, 
with aeropyles absent. Micropylar plug at Exochorion ca. 4 X thicker than endo- 

anterior pole, ovate, and with two distinct, chorion. Pore canals widest at base 

lateral helical micropylar tubes (Fig. 7). (Fig. 9). 



VOLUME 101. NUMBER 1 








VV)! 



JgS^Hf^ 


^BIBI^^ IIM^pi^Hbr 2i%SI3l9 


' '^'V ' '- 


u 






r r'«T*^: isg^" ;« 




11 H 






Figs. 8-13. 8. 9, Ambry sus circunicinctus. 8, Micropyle section at base of plug. 9, Chorion section. 10, 11, 
A. montandoni. 10, Anterior pole with micropylar plug. 1 1, Chorion surface. 12, 13, A. arizonus. chorion surface. 



Ambrysus (SyncoUus) montandoni La 

Rivers 

(Figs. 10, 11) 

Ambrysus montandoni La Rivers 1963: 1—5. 
VENEZUELA: Territorio Amazonas, To- 
bogan 



Length, 1.34; width, 0.64; n = 1; dissected. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of te- 
tragonal to heptagonal units, delimited by 
sulci between adjacent hemispherical, fu- 
runcular mounds (Fig. 10). Within each 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



unit, 15-30 aeropyles, becoming less dis- 
tinct toward anterior pole. Aeropyles in sul- 
ci as well as on mounds. Numerous glob- 
ules adhered to surface (Fig. 1 1). (Although 
these globules may be artifacts, they were 
persistent even after sonication in 100% 
acetic acid.) Reticulation less conspicuous 
anteriorly because mounds become flat- 
tened until immediate vicinity of micropyle 
where smaller reticulation units are evident 
(Fig. 10). Micropylar plug at anterior pole 
and amorphous. Protuberances other than 
micropyle, mounds, and granules lacking. 

Ambrysus (Ambrysus) arizonus La Rivers 
(Figs. 12-15) 

Ambrysus arizonus La Rivers 1951: 320— 

322. 
USA; Arizona, Gila Co., Jakes Corner 
Length, 1.45 ± 0.01; width, 0.77 ± 0.01; n 

= 8; oviposited. 

Overall appearance elongate-oval. Retic- 
ulation generally consisting of pentagonal 
to heptagonal units, delimited by depres- 
sions. Each depression with double row of 
elongate papillae defining unit boundaries 
(Fig. 12). Within each unit, chorion raised 
and coarsely papillose (Fig. 13). Aeropyles 
indistinct, evident as pitted appearance 
among papillae; number 20-50 per cell. Tu- 
bercles lacking. Anterior pole with reticu- 
lation and papillae less distinct. Micropylar 
plug amorphous; section through base of 
micropylar plug with two micropyles (Fig. 
14). 

Exochorion distinctly thicker than endo- 
chorion. Pore canals widest in basal third 
(Fig. 15). 

Ambrysus (Ambrysus) buenoi Usinger 
(Figs. 16, 17) 

Ambrysus buenoi Usinger 1946: 199-200. 
USA: Texas, Kimble Co., Junction 
Length, 1.32 ± 0.02; width, 0.65 ± 0.02; n 
= 11; oviposited. 

Overall appearance elongate-oval. Sur- 
face comprising a series of anastomosing 
mounds with irregularly produced protuber- 



ances (Fig. 16). Bases of mounds with large 
aeropyles distributed randomly. Chorionic 
surface, including swells and protuberanc- 
es, granular (Fig. 17). Micropyle amor- 
phous. Chorionic surface immediately sur- 
rounding micropyle lacking regular surface 
features, although poorly defined mounds 
may occur. 

Ambrysus (Ambrysus) crenulatus 

Montandon 

(Figs. 18, 19) 

Ambrysus crenulatus Montandon 1897a: 

13-14. 
ECUADOR: Napo Province, Puerto Napo 
Length, 1.12 ± 0.01; width, 0.49 ± 0.01; n 

= 10; dissected. 

Overall appearance elongate-oval. Retic- 
ulation generally consisting of pentagonal 
to heptagonal units, delimited by distinctly 
raised, thin walls (Fig. 18). Each unit ap- 
pearing as a deep socket, with approxi- 
mately 8-17 large, irregularly distributed 
aeropyles (Fig. 19). Tubercles lacking. An- 
terior pole with reticulation less distinct. 
Micropylar plug amorphous. Protuberances 
other than micropyle and raised reticulation 
lacking. 

Ambrysus (Ambrysus) fossatus Usinger 
(Figs. 20, 21) 

Ambrysus fossatus Usinger 1946: 191-192. 
ECUADOR: Napo Province, Puerto Napo 
Length, 1.23 ± 0.03; width, 0.57 ± 0.02; n 
= 3; dissected. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of pen- 
tagonal to heptagonal units and only faintly 
visible (Fig. 20). Within each unit, 10-20 
conspicuous aeropyles. Chorionic surface 
devoid of protruberances other than micro- 
pyle. Middle of pore canals generally par- 
allel-sided (Fig. 21). Micropylar plug amor- 
phous, acentric. 

Ambrysus (Ambrysus) hungerfordi Usinger 
(Figs. 22-25) 

Ambrysus hungerfordi Usinger 1946: 192- 
194. 



VOLUME 101. NUMBER 1 




^^^-_-» - • -t 



M'^7*iiisii:M^'. 




Figs. 14-19. 14. 15. Ambrysus arizotnis. 14, micropyle section at base of plug. 15. Chorion section. 16, 17. 
A. buenoi. chorion surface. 18, 19. A. ciemilatiis, chorion surface. 



USA: Texas, Presidio Co., Big Bend Ranch Overall appearance oval (Fig. 22). Retic- 
State Natural Area ulation pattern generally consisting of pen- 
Length, 1.18 ± 0.01; width, 0.63 ± 0.01; n tagonal to heptagonal units, delimited by 
= 9; dissected. low, raised ridges (Fig. 23). Within each 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 








Figs. 20-25. 20, 21, Ambrysus fossatus. 20, Chorion surface. 2L Chorion section. 22-25, A. Iningeiforcli. 
22, Whole egg. 23, Chorion surface. 24, Micropylar plug. 25, Chorion section. 



unit, 50-120 aeropyles more or less evenly 
distributed. Protruberances, tubercles, and 
papillae lacking. Micropylar plug anterior 
and amorphous (Fig. 24). Anterior pole 



with reticulation less pronounced, aeropyles 
fewer. Chorionic inner layer 60% as thick 
as chorionic outer layer (Fig. 25). Pore ca- 
nals narrowest at middle. 



VOLUME 101. NUMBER 1 




Figs. 26-31. 26, Ambrysus inflatus. chorion surface. 27-29, A. lunatus. 11, Chorion section. 28, Pore canal 
containing bacteria. 29, Micropyle .section at base of plug. 30, 31, y4. mormon, chorion surface. 



Ambrysus (Ambrysus) inflatus La Rivers 
(Fig. 26) 

Ambrysus inflatus La Rivers 1953a: 1316- 

1318. 
MEXICO: Jalisco, Chapala 



Length, 1.37 ±0.01; width, 0.68 ± 0.01; n 
= 10; dissected. 

Overall appearance elongate with 
rounded ends. Reticulation pattern gener- 
ally consisting of pentagonal to heptago- 



10 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 32-37. 32, Ainbiysiis mormon, anterior pole with micropylar plug. 33. A. occidentalis. chorion .surface 
with anthefomi processes. 34, 35, A. phiutus. chorion surface. 36. 37, A. portheo. 36, Anterior pole. 37, Chorion 
surface with antheform processes. 



nal units, delimited by a series of irregu- 
larly raised ridges with numerous tiny pa- 
pillae between (Fig. 26). Scattered groups 
of papillae depressed, creating pitted ap- 
pearance. Anterior pole with amorphous 



micropylar plug, without raised reticula- 
tion or papillae. 

The appearance of the chorion is virtu- 
ally indistinguishable from that of A. luna- 
tus Usinger (see Sites and Nichols 1990). 



VOLUME 101, NUMBER 1 



11 



Ambrysus (Ambrysus) limatus Usinger 
(Figs. 27-29) 

Ambrysus limatus Usinger 1946: 203-205. 
Ambrysus lunatus: Sites and Nichols 1990: 

800-808. 
USA: Texas, Kimble Co., Junction 

Endochorion subequal to exochorion in 
thickness (Fig. 27). Pore canals parallel-sid- 
ed and widest at base. Presence of bacteria 
in pore canals as detected by transmission 
electron microscopy (Fig. 28). Micropylar 
plug with three micropyles (Fig. 29). 

Original description of egg was given by 
Sites and Nichols (1990). 

Ambrysus (Ambrysus) mormon Montandon 
(Figs. 30-32) 

Ambrysus monnon Montandon 1909: 48-49. 
Ambrysus mormon: Usinger 1946: 186— 

187, Plate X. 
USA: New Mexico, Lincoln Co., Hondo 
Length, 1.70 ± 0.03; width, 0.98 ± 0.02; n 

= 6; oviposited. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of te- 
tragonal to heptagonal units, delimited by a 
series of depressions between composite tu- 
mescences (Fig. 30). Tumescences rarely 
entire, usually completely or incompletely 
divided into two to four components, oc- 
casionally with a smaller, central tumes- 
cence or depression (Fig. 31). Tumescences, 
depressions, including reticulation, covered 
with papillae. Aeropyles generally distrib- 
uted, neither clustered nor concentrated. Tu- 
mescences becoming flatter and less divid- 
ed toward anterior pole. Micropylar plug 
amorphous, acentric (Fig. 32). 

Usinger (1946) indicated that eggs are 
glued to the surface of pebbles and are sub- 
oval with a buttonlike micropyle at the an- 
terior pole. 

Ambrysus (Ambrysus) occidentalis La 
Rivers 

(Fig. 33) 

Ambrysus occidentalis La Rivers 1951: 
322-325. 



USA: Arizona, Gila Co., Jakes Comer 
Length, 1.28 ± 0.01; width, 0.62 ± 0.00; n 
= 3; dissected. 

Overall appearance elongate-oval. Reticu- 
lation pattern generally consisting of hexag- 
onal units, delimited by raised boundaries. 
Single antheform process extending outward 
from center of each unit, distal end concave 
and expanded (Fig. 33). Margins surrounding 
distal concavity irregular, never in contact 
with adjacent antheform process. Base of an- 
theform process widest, gradually narrowing 
distally. Papillae covering surface from raised 
reticulation to base of antheform process. 25- 
40 irregularly distributed aeropyles distinctly 
visible around base of antheform process. 
Anterior pole with micropylar plug, with pat- 
tern less distinct, antheform processes and pa- 
pillae absent. 

Ambrysus (Ambrysus) plautus Polhemus 
and Polhemus 
(Figs. 34, 35) 

Ambrysus plautus Polhemus and Polhemus 

1982: 326-328. 
MEXICO: Chihuahua, Cusarare 
Length, 1.12 ± 0.01; width, 0.54 ± 0.02; n 

= 3; dissected. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of pen- 
tagonal to heptagonal units, delimited by 
depressed lines. Within each unit, chorion 
raised and distinctly papillose (Fig. 34). 
Aeropyles large, distinct; number 3-12 in 
interior of unit, 10-18 in depressed perim- 
eter of unit (Fig. 35). Tubercles lacking. 
Anterior pole with pattern and papillae less 
distinct. Micropylar plug amorphous. 

Ambrysus (Ambrysus) portheo La Rivers 
(Figs. 36-39) 

Ambrysus portheo La Rivers 1953a: 1320- 

1321. 
MEXICO: Nuevo Leon, La Nogalera 
Length, 1.56 ± 0.02; width, 0.89 ± 0.01; n 

= 2; oviposited. 



12 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 38-43. 3S. 3M, Xnihiysiis portheo. 38, Micropyle section at base ot plug. 39, Chorion section. 40-43, 
A. pudiciis. 40, Anterior pole with micropylar plug (dislodged). 41, Chorion surface. 42, Micropyle section at 
base of plug. 43, Chorion section. 



General appearance robust with apices 
slightly truncate (Fig. 36). Reticulation 
pattern generally consisting of pentagonal 
to hexagonal units, delimited by raised. 



boundary with irregularly-spaced, deep 
notches extending ca. y, distance to base. 
Single antheform process extending out- 
ward from center of each unit, the distal 



fence-like boundaries. Ectal edge of end of which is concave and expanded 



VOLUME 101, NUMBER 1 



13 



(Fig. 37). Margins surrounding distal con- 
cavity irregular, never in contact with ad- 
jacent antheform process. Base of anthe- 
form process widest, gradually narrowing 
distally. Papillae covering surface be- 
tween unit boundaries and base of anthe- 
form process. Irregularly spaced aeropy- 
les occasionally visible in gaps between 
papillae. Anterior pole with pattern re- 
duced around micropylar plug, antheform 
processes and papillae absent. Micropylar 
plug with two micropyles (Fig. 38) 

Exochorion slightly thicker than endo- 
chorion. Pore canals bulbous at base. An- 
theform processes solid, without ducts (Fig. 
39). 

Ambt^'sus (Ambrysus) pudicus Stal 
(Figs. 40-43) 

Ambrysus pudicus Stal 1862: 460. 
USA: Texas, Kimble Co., Junction 
Length, 1.12 ± 0.01; width, 0.61 ± 0.01; n 
= 11; oviposited. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of pen- 
tagonal to heptagonal units, delimited by 
slightly raised boundaries (Fig. 40). Within 
each unit, protuberances of two sizes: larger 
tubercles and smaller pustules (Fig. 41). Pe- 
rimeter of each unit with a series of ca. 12- 
25 pustules; additional pustules usually near 
center of unit and among tubercles. Tuber- 
cles number 1-5 per unit and situated near 
center, occasionally surrounding one or 
more pustules. Outline of pustules round, of 
tubercles amoebiform. Anterior pole with 
pattern faintly visible, protuberances other 
than micropyle lacking. Micropylar plug 
amorphous, acentric, with two micropyles 
(Fig. 42). 

Exochorion thicker than endochorion 
(Fig. 43). Pore canals wide, ca. 0.4 X 
length. 

Ambrysus (Ambrysus) pulchellus 

Montandon 

(Figs. 44, 45) 

Ambrysus pulchellus Nioniandon 1897a: 16. 
USA: Texas, Kimble Co., Junction 



Length, 1.36 ± 0.02; width, 0.65 ± 0.01; n 
= 11; oviposited. 

General appearance elongate-oval with 
rounded apices. Reticulation pattern gener- 
ally consisting of pentagonal to heptagonal 
units, delimited by sulci between tumes- 
cences (Fig. 44). Each tumescence extends 
around perimeter of unit and abuts adjacent 
tumescences (Fig. 45). Single, smaller, ir- 
regularly-shaped tumescence within each 
perimeter tumescence. Occasionally, central 
tumescence absent, replaced by depression 
near center. Tumescences generally gla- 
brous. Clusters of approximately 12-20 
aeropyles distributed over surface of tu- 
mescences, concentrated near margins. Mi- 
cropylar plug slightly acentric and amor- 
phous. 

Ambrysus (Ambrysus) puncticollis Stal 
(Figs. 46-48) 

Ambrysus puncticollis Stal 1876: 143. 
USA: Texas, Kimble Co., Junction 
Length, 1.36 ± 0.01; width, 0.66 ± 0.01; n 
= 11; oviposited. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of pen- 
tagonal to heptagonal units, delimited by 
depressed lines (Fig. 46). Within each unit, 
20-80 large aeropyles (aeropyles filled with 
debris in SEMs) (Fig. 47). Protruberances, 
tubercles, and papillae lacking. Micropylar 
plug anterior and amorphous. Anterior pole 
with pattern less distinct. 

Exochorion ca. 4.5 X thicker than endo- 
chorion (Fig. 48). Pore canals slightly di- 
vergent entally and occur at somewhat reg- 
ular interval. 

Ambrysus (Ambrysus) spiculus Polhemus 
and Polhemus 
(Figs. 49-51) 

Ambrysus spiculus Polhemus and Polhemus 

1981: 400-401. 
MEXICO: Chihuahua, Rio Concheiio 
Length, 1.10 ± 0.01; width, 0.48 ± 0.03; n 

= 3; dissected. 



14 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 44-49. 44, 45, Ambiysits piilchelhis. chorion surface. 46-48. A. piincticollis. 46, 47, Chorion surface. 
48, Chorion section. 49, A. spiciiliis. whole egg. 



Overall appearance elongate-oval (Fig. 
49). Reticulation pattern generally consist- 
ing of pentagonal to heptagonal units, de- 
limited by depressed boundaries. Within 
each unit, chorion raised and finely papil- 



lose (Fig. 50). Perimeter of each unit with 
8-16 large, distinct aeropyles; near the cen- 
ter of unit 1-4 aeropyles. Anterior pole 
with reticulation and papillae less evident. 
Micropylar plug amorphous (Fig. 51). 



VOLUME 101. NUMBER 1 



15 




Vvji*:-? 3^^: 









Figs. 50-55. 50. 51. Amhrysiis spiciilus. 50. Chorion surface. 51. Anterior pole with micropylar plug. 52- 
54. A. thermaruin. 52. 53. Chorion surface. 54. Chorion section. 55, A. triilenlatits. chorion surface. 



Ambrysus (Ambrysus) thermarum La Length, L43 ± 0.06; width, 0.72 ± 0.02; n 

Rivers = 10; oviposited. 

igs. J -J ) Overall appearance elongate-oval. Retic- 
Ambrysus thennarum La Rivers 1953b: 1-3. ulation pattern generally consisting of pen- 
USA: New Mexico, Taos Co., Arroyo Hondo tagonal to heptagonal units, delimited by 



16 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 56-6 L 56, Ambi-ysiis tridentatus, chorion surface. 57, 58, A. woodburyi. chorion surface. 59-61, 
Gestroiella liiiiiiocoroides. 59, Whole egg. 60, Chorion surface. 61. Micropylar plug. 



double row of rounded to elongate papillae cence (Fig. 53). Aeropyles distinct and ran- 

(Fig. 52). Chorionic surface generally cov- domly distributed on chorion except on tu- 

ered with papillae. Within each unit formed mescence; number 15-30 per unit. Pattern, 

by reticulation, a single amorphous tumes- tumescences, papillae, aeropyles becoming 



VOLUME 101. NUMBER 1 



17 



less distinct toward anterior pole. Micro- 
pylar plug amorphous. 

Exochorion slightly thicker than endo- 
chorion (Fig. 54). Pore canals widest at 
base. 

Ambrysus (Ambrysus) tridentatus La 

Rivers 

(Figs. 55, 56) 

Ambrysus tridentata La Rivers 1962: 129- 

132. 
MEXICO: Nuevo Leon, Potrero Redondo 
Length, 1.14; width, 0.58; n = 1; dissected. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally consisting of pen- 
tagonal to heptagonal units, delimited by 
depressed boundaries. Chorion domed 
within each unit (Fig. 55). Aeropyles sub- 
circular near center of dome, ellipsoid off- 
center; 20-40 per cell. Chorion smooth; pa- 
pillae and tubercles lacking (Fig. 56). An- 
terior pole with domes diminishing in size, 
aeropyles becoming less evident. Micropy- 
lar plug amorphous. 

Ambrysus (Ambrysus) woodburyi Usinger 

(Figs. 57, 58) 
Ambrysus woodburyi Usinger 1946: 194- 

195. 
USA: Arizona, Cochise Co., Portal 
Length, 1.16 ± 0.02; width, 0.57 ± 0.02; n 
= 10; oviposited. 

Overall appearance elongate-oval. Retic- 
ulation pattern generally poorly-defined. 
Within each unit formed by reticulation, a 
single amorphous tumescence (Fig. 57). 
Each unit with approximately 7-12 large, 
distinct aeropyles randomly distributed 
around the tumescence. Chorionic surface 
generally finely papillose (Fig. 58). Ante- 
rior pole with tumescences and papillae less 
developed. Micropylar plug amorphous, 
acentric. 

Subfamily Cheirochelinae Montandon 

1897b 

Genus Gestroiella Montandon 1897b 

Gestroiella limnocoroides Montandon 

(Figs. 59-61) 

Gestroiella limnocoroides Montandon 

1897b: 371-372. 



THAILAND: Chiang Mai Prov., Chiang 

Mai 
Length, 3.04 ± 0.02; width, 1.35 ± 0.02; n 

= 4; dissected. 

Overall appearance elliptical with poles 
slightly acuminate (Fig. 59). Reticulation 
pattern generally consisting of pentagonal 
to octagonal units, delimited by low, ele- 
vated ridges (Fig. 60). Within each unit, 
300-800 aeropyles. Micropyles incorpo- 
rated into low, broad mound at anterior 
pole (Fig. 61). Other than micropyle, pro- 
truberances, tubercles, and papillae lack- 
ing. 

Subfamily Aphelocheirinae Fieber 1851 

Genus Aphelocheirus Westwood 1833 

Aphelocheirus femoratus Polhemus and 

Polhemus 

(Figs. 62-64) 

Aphelocheirus femoratus Polhemus and 

Polhemus 1988: 214-216. 
THAILAND: Songkhla Prov., Ton Nga 

Chang National Park 
Length, 1.13 ± 0.01; width, 0.51 ± 0.01; n 

= 10; dissected. 

General appearance oval and robust with 
anterior pole slightly truncate (Fig. 62). Re- 
ticulation pattern generally consisting of 
pentagonal to heptagonal units, delimited 
by broad, slightly elevated ridges (Fig. 63). 
Within each unit, 100-300 aeropyles. Mi- 
cropyle amorphous (Fig. 64). Chorionic 
surface immediately surrounding micropyle 
lacking well-defined surface features. Other 
than micropyle, protruberances, tubercles, 
and papillae lacking. 

Subfamily Laccocorinae Stal 1876 

Genus Heleocoris Stal 1876 

Heleocoris ovatus Montandon 

(Figs. 65-67) 

Heleocoris ovatus Montandon 1897c: 451- 

452. 
THAILAND: Yala Province, Than To, 

Banglang National Park 
Length, 1.55 ± 0.01; width, 0.71 ± 0.01; n 

= 9; dissected. 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





o >■ _ :, . 


^ ■'" ' "''i--- "'-"i! 


P .■:- 


, .:.-_ '.\'^^^ 


; ..-'". 


-•*"ft*>«^ 


-- ' "-'"'' •'' '-jF' 


. '^- : . 


-" " -'l ■'■ ■-- -,' •'', 


i"-'/-''' • ' 


i *— "jlrj^* 




finawrli. 


iii,^- 


^-k;.- - 




^- 


. " , !• 


V.-';-. ^. _■ 


- . , j' 


'--- . 


^^^M.:i«><^" . 


' :«:' 


': — ,^, ' - • 


«<arfi»*^ 


63 1 


20n , I 






'-'*— ^ ' ' ^^"!58II^^BKHBr 




Figs. 62-67. 62-64. Aphelocheirus femoratits. 62, Whole egg. 63. Chorion surface. 64, Anterior pole with 
micropylar plug. 65-67, Heleocoris ovatiis. 65, Whole egg. 66, Chorion surface. 67, Micropylar plug in concavity 
at anterior pole. 



Overall appearance elongate, parallel- 
sided with rounded ends (Fig. 65). Reticu- 
lation pattern generally consisting of tetrag- 
onal to heptagonal units, delimited by cos- 



tiform ridges (Fig. 66). Within each unit, 
25-75 aeropyles. Chorionic surface gener- 
ally granular. Anterior pole with amorphous 
micropylar plug set in acetabular depression 



VOLUME 101. NUMBER 1 



19 




liiil 


KB 


^SH 




« 
* 




«gbeg 


<• 




^1 






^•ijfl 



^ 


p 


PI 






^BI^W| 


HlJS^ 












R 




■ 


ImBR^^ 


^^^K'* 


^. 


--■^ » 












* 


■■■?'i. 




HP^f*' 




1 


Ih^'^ 


di 


ri 


■ 


ni' !» '' ^1 


■ 


^^^^^^■i^H 


H^H 


rtgJM 


^H 


9 


■ 




■ 




Figs. 68-73. 68-71, Lininocoris moapensis. 68. Whole egg. 69. 70. Micropyles. 71. Chorion section. 72. 
73, Ilyocoris cimicoides. 72. Whole egg. 73. Anterior pole. 



(Fig. 67). Pattern in depression becoming tus. Locality and ecological data associated 

indistinct toward micropyle. with the collection of the adults from which 

Pending taxonomic revision, this species eggs were obtained are available in Sites et 

has been determined probably to be H. ova- al. (1997). 



20 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Subfamily Limnocorinae Stal 1876 

Genus Limnocoris Stal 1858 

Limnocoris moapensis (La Rivers) 

(Figs. 68-71) 

Usiiigerina moapensis La Rivers 1950: 

368-373. 
Limnocoris moapensis: Sites and Willig 

1994: 810. 
USA: Nevada, Clark Co., Moapa 
Length, 0.98 ± 0.01: width, 0.56 ± 0.01; n 

= 8; oviposited. 

Overall appearance robust, elongate-oval 
(Fig. 68). Reticulation pattern generally con- 
sisting of pentagonal to heptagonal units, de- 
limited by faintly impressed lines. Within 
each unit, chorion smooth, perforated by 1 5- 
60 aeropyles set in shallow individual sock- 
ets. Chorion smooth, devoid of papillae, tu- 
bercles, and protruberances other than mi- 
cropyle. Anterior pole with micropyle set in 
concavity, immediately surrounded by mod- 
ified chorion devoid of pattern and aeropy- 
les. 2-3 distinct micropyles (v = 2.8, n = 
10) arising from central point, extending 
outward in arcuate fashion (Figs. 69, 70). 

Exochorion ca. % as thick as endocho- 
rion. Pore canals widest at base (Fig. 71). 

Subfamily Naucorinae Stal 1876 

Genus Ilyocohs Stal 1861 

Ilyocoris cimicoides (Linnaeus) 

(Figs. 72-77) 

Nepa cimicoides Linnaeus 1758: 440. 
Ilyocoris cimicoides: Stal 1861: 201. 
Naucoris cimicoides: Rawat 1939: 123— 

127, Figs. 1-3. 
CZECHOSLOVAKIA: southern Bohemia, 

Veselnad Lu Nic 
Length, 2.25 ± 0.05; width, 0.56 ± 0.03; n 

= 8; dissected. 

Overall appearance cylindrical and elon- 
gate with a 45 degree bend near middle 
(Fig. 72). Anterior pole flattened (Fig. 73), 
posterior pole rounded. Reticulation gener- 
ally consisting of pentagonal to heptagonal 
units, delimited by faintly impressed 
boundaries (Fig. 74). Within each unit, cho- 



rion smooth, perforated by 50-90 aero- 
pyles. Flattened anterior pole with elongate 
tumescences radiating outward from acen- 
tric micropylar plug (Fig. 75). Micropylar 
plug with four micropyles (Fig. 76), each 
of which is raised slightly above the re- 
mainder of the plug (Fig. 77). 

Rawat (1939) described the egg as ap- 
proximately 2 mm in length and cylindrical 
with an operculate, recurved end. Lebrun 
(1960) illustrated the position of the micro- 
pylar plug on the operculum (although not 
labeled as such) and Hinton ( 1981) reported 
the presence of three to four micropyles. 
This is the only naucorid documented to 
have endophytic oviposition (Rawat 1939, 
Cobben 1968). 

Genus Pelocoris Stal 1876 

Pelocoris femoratus (Palisot de Beauvois) 

(Figs. 78-81) 

Naucoris femorata Palisot de Beauvois 

1820: 237. 
Pelocoris femoratus: Stal 1876: 144. 
Pelocoris femorata: Torre Bueno 1903: 

168-172. 
Pelocoris carolinensis: Hungerford 1927: 

80-82, Plate VI. 
Pelocoris femoratus: McPherson et al. 

1987: 291. 
USA: Missouri, Boone Co., Columbia 
Length, 1.17 ± 0.01; width, 0.64 ± 0.01; n 

= 10; oviposited. 

Overall appearance elongate-oval (Fig. 
78). Reticulation pattern generally consisting 
of pentagonal to heptagonal units. Within 
each unit, irregular and sometimes discontin- 
uous elongate tumescence approximating 
boundary (Fig. 79). Anterior pole with mi- 
cropylar plug set in shallow concavity. Mi- 
cropylar plug somewhat amorphous and in- 
consistent in appearance, with micropyles 
opening laterally (Fig. 80). Number of mi- 
cropyles indistinct, but apparently 2—3. 

Exochorion ca. 3.5 X thicker than en- 
dochorion. Pore canals widest at middle and 
base (Fig. 81). 

Using light microscopy, McPherson et 



VOLUME 101, NUMBER 



21 





n "^:^ " ^**ugr u mil, '"■•o^^ 


sk 


i^l^^^llllf'!^]^^^ y^^^^Lylj 


W^ 


^^^^ 


1 


^^^ 


fc 

N 


7y^-H'-.^<«^',^,^- 



Figs. 74-79. 74-77, llyocoris cimicoides. 74. Chorion surface. 75, Anterior pole. 76, Micropylar plug. 77. 
Micropylar opening. 78. 79, Pelocohs femoratiis. 78, Whole egg. 79. Chorion surface. 



al. (1987) reported that the chorion has a Discussion 

primarily irregular hexagonal pattern and 

the micropylar plug is situated at the ante- Differences were evident in the chorion 

rior end. between incompletely developed and well- 



22 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 80-81. Pelocoris femoratus. 80, Micropylar plug. 8L Chorion section. 



developed eggs. Eggs that were incomplete- 
ly developed generally appeared to have a 
single point within each reticulation unit, 
which was raised and around which the 
chorion appeared to have flowed down onto 
the surface of the egg. This 'poured' ap- 
pearance probably represented the site of 
chorion deposition for each follicular epi- 
thelial cell. Nonetheless, for dissected eggs, 
even though we selected eggs from the 
common oviduct or vagina for descriptions, 
the possibility exists that egg structure may 
have continued to develop prior to ovipo- 
sition. For the 21 species of Ambtysus ex- 
amined, chorionic sculpturing differs inter- 
specifically and generally is species-specif- 
ic. Although these differences were noted, 
other specific features were common among 
some of the species. 

Previously, eggs were described and 
electron micrographs presented for A. lun- 
atus (Sites and Nichols 1990), a member of 
the signoreti group. Other members of the 
signoreti group represented here are A. in- 
flatiis, A. occidentalis, and A. portheo. Gen- 
erally, these four species share egg features 
including an acutely raised, fence-like retic- 
ulation forming a polygonal pattern, and 
minute papillae distributed generally over 
the surface. In addition, two species possess 
elongate, antheform processes. Eggs of oth- 
er members of the signoreti group are likely 
to possess these features. 



Eggs of the genus Ambrysus have been 
reported to have six micropyles (Hinton 
1981), and those of an unspecified species 
from Aruba, Netherlands Antilles, usually 
have at least five (Cobben 1968). The in- 
dividual micropylar tubes of Ambrysus are 
fused into a single, prominent plug (Cobben 
1968). Our internal examinations have re- 
vealed two micropyles in each of three spe- 
cies of Ambrysus and three in two other 
species. Although it is likely that intraspe- 
cific variation exists in micropyle number 
for species of Ambrysus, as has been ob- 
served in species of other naucorid genera 
[e.g., C. hungerfordi (Sites and Nichols 
1993)], we have observed only two and 
three micropyles. Thus, our data do not cor- 
roborate Hinton's (1981) report of six nor 
Cobben's (1968) report of five or more mi- 
cropyles for species of Ambrysus. 

In sharp contrast to Ambrysus, the num- 
ber of micropyles for species of Limnocoris 
is clear with external examination because 
micropylar fusion is minimal. For Limno- 
coris lutzi and Limnocoris sp. [Ecuador, see 
Sites (1990)], two micropyles are clearly 
evident. Of 10 eggs of Limnocoris moap- 
ensis (La Rivers), eight had three micro- 
pyles whereas the other two had two mi- 
cropyles. Previous reports for Limnocoris 
micropyles are nonexistent. 

The number of micropyles for species of 
Pelocoris is unclear and the degree of fu- 



VOLUME 101. NUMBER 1 



23 



sion differs interspecifically. Sites (1991) 
revealed two partially-fused micropyles for 
P. poeyi. Surprisingly, for P. femora tus, the 
micropyle number has not previously been 
given despite three separate descriptions of 
eggs [Torre Bueno 1903, Hungerford 1927 
as P. carolinensis (see La Rivers 1948b), 
McPherson et al. 1987]. Pelocoris femora- 
tiis micropyles are fused into a plug ['mi- 
cropylar boss' of Torre Bueno (1903)] sim- 
ilar to that of Ambrysus. The form of the 
plug is inconsistent, and a canal leading to 
a micropylar opening may be observed in 
some specimens. 

In the only report for eggs of species of 
the subfamily Potamocorinae, which is con- 
sidered by some to represent a distinct fam- 
ily level taxon (e.g., Stys and Jansson 
1988), Cobben (1968) indicated that C. kle- 
erekoperi has a single micropylar opening 
with several external mucous projections. 

Systematic Value 

The family Naucoridae is not blessed 
with even a modicum of somatic characters 
that varies among the higher taxa that may 
be used to elucidate systematic relation- 
ships. The principal characters that have 
been used for interspecific taxonomic dis- 
tinctions have been adult male and female 
external genitalic features. Characters of 
nymphs and eggs have not been used, al- 
though Lopez Ruf (1989) suggested that 
chorionic attributes may be valuable taxo- 
nomic characters externally at the species 
level and internally at the generic level. We 
concur with this assessment. Specifically, 
intergenerically variable characters include 
the relative widths of the chorionic inner 
and outer layers and pore canal configura- 
tion. Although the number of micropyles 
does not vary appreciably among these gen- 
era, the degree of external fusion of the in- 
dividual micropylar tubes may be a taxo- 
nomically valuable character at generic or 
higher levels. External chorionic patterns 
are quite valuable as an interspecific diag- 
nostic character in certain genera (e.g., Am- 
brysus). However, the pattern is invariant 



among the four species of Limnocoris that 
we have examined. Thus, the utility of this 
character in providing systematic resolution 
appears to be restricted to particular genera. 

Acknowledgments 

We thank D. L. Pinkerton and J. A. 
White, Electron Microscopy Facility, Col- 
lege of Agriculture, University of Missouri, 
for assistance with the electron microscopy 
and preparation of micrographs; and J. T 
Polhemus and Jitka Vilimnova for loan or 
gift of selected females. We also thank T. J. 
Henry, J. E. McPherson, and J. T. Polhemus 
for critical reviews of this manuscript, and 
T. J. Henry for advice on nomenclature. 
Funding for RWS was provided in part by 
MU project #PSSL0232. This is Missouri 
Agricultural Experiment Station journal se- 
ries paper No. 12,744. 

Literature Cited 

Clarke. E and E. Baroudy. 1990. Studies on Lacco- 
coiis limigeniis (Stal) (Hemiptera: Naucoridae) in 
Lake Naivasha, Kenya. The Entomologist 109: 
240-249. 

Cobben, R. H. 1968. Evolutionary trends in Heterop- 
tera: Part I: Eggs, architecture of the shell, gross 
embryology and eclosion. Centre for Agricultural 
Publishing and Documentation, Waginengen. 

Eieber, E X. 1851. Genera Hydrocoridum secundum 
ordinem naturalem in familias disposita. Abhand- 
lung der Koniglische Bohemischen gesellschaft 
der Wissenschaften in Prag, Series 5, 7: 181-212, 
4 plates. [Separate: Actis Regiae Behemicae So- 
cietatis Scientarium, Pragae, 1: 130, 4 plates]. 

Hinton, H. E. 1981. Biology of insect eggs (3 vols.). 
Pergamon Press, Oxford. 

Hungerford. H. B. 1927. The life history of the creep- 
ing water bug, Pelocoris carolinensis Bueno 
(Naucoridae). Bulletin of the Brooklyn Entomo- 
logical Society 22: 77-83. 

La Rivers. I. 1948a. A new species oi Amh)-\siis from 
Death Valley, with notes on the genus in the Unit- 
ed States (Hemiptera: Naucoridae). Bulletin of the 
Southern California Academy of Sciences 47: 
103-110. 

. 1948b. A new species of Pelocoris from Ne- 
vada, with notes on the genus in the United States 
(Hemiptera: Naucoridae). Annals of the Entomo- 
logical Society of America 41: 371-376. 

. 1950. A new naucorid genus and species from 

Nevada (Hemiptera). Annals of the Entomological 
Society of America 43: 368-373. 



24 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



. 195L A revision of the genus Ambnsus in 

the United States (Hemiptera: Naucoridae). Uni- 
versity of CaHfornia Publications in Entomology 
8: 277-338. 

. 1953a. The Ainhiysus of Mexico (Hemiptera, 

Naucoridae). The University of Kansas Science 
Bulletin 35: 1279-1349. 

. 1953b. Two new naucorid bugs of the genus 

Ambi-ysus. Proceedings of the United States Na- 
tional Museum 103: 1-7. 

. 1962. A new species of Ambrysiis from Mex- 
ico (Hemiptera: Naucoridae). The Wasmann Jour- 
nal of Biology 20: 129-133. 

. 1963. Two new Ambrysi (Hemiptera: Nau- 
coridae). Biological Society of Nevada Occasional 
Papers 1: 1-7. 

. 1971. Studies of Naucoridae (Hemiptera). Bi- 
ological Society of Nevada Memoirs 2, iii -I- 120 
pp. 

. 1974. Catalogue of taxa described in the fam- 
ily Naucoridae (Hemiptera) supplement no. 1: 
Corrections, emendations and additions, with de- 
scriptions of new species. Biological Society of 
Nevada Occasional Papers 38: 1-17. 

. 1976. Supplement no. 2 to the catalogue of 



taxa described in the family Naucoridae (Hemip- 
tera), with descriptions of new species. Biological 
Society of Nevada Occasional Papers 41: 1-17. 

Larsen, O. 1927. Uber die Entwicklung und Biologic 
von Aphelocheirus cpstivalis Fabr. Entomologisk 
Tidskrift 48: 180-206. 

Lebrun, D. 1960. Recherches sur la biologic et 
I'ethologie de quelques Heteropteres aquatiques. 
Annales de la Societe Entomologique du France 
129: 179-199. 

Linnaeus, C. 1758. Systema Naturae, 10th ed. Hol- 
miae, 823 pp. 

Liu, G. Q. and L. Y. Zheng. 1994. Two new species 
of genus Aphelocheirus Westw. from China (In- 
secta: Heteroptera: Aphelocheiridae). Reichenba- 
chia 30: 113-117. 

Lopez Ruf, M. L. 1989. Los huevos de algunas espe- 
cies de los generos Pelocoris y Ainhnsiis (Het- 
eroptera Limnocoridae). Limnobios 2: 720-724. 

McPherson, J. E., R. J. Packauskas, and P. P. Korch, 
III. 1987. Life history and laboratory rearing of 
Pelocoris femoratus (Hemiptera: Naucoridae), 
with descriptions of immature stages. Proceedings 
of the Entomological Society of Washington 89: 
288-295. 

Montandon, A. L. 1897a. Hemiptera Cryptocerata. 
Earn. Naucoridae. -Sous. fam. Cryptocricinae. Ver- 
handlungen der Kaiserlich-Koniglichen Zoologis- 
che-Botanischen Gesellschaft in Wien 47: 6-23. 

. 1897b. Viaggio di Leonardo Fea in Birmania 

e Regioni Vicine. LXXV. Hemiptera Cryptocerata. 
Annali del Museo Civico di Storia Naturale di 
Genova 17(2): 365-377. 



. 1897c. Hemiptera Cryptocerata. Fam. Nau- 
coridae. -Sous-fam. Laccocorinae. Verhandlungen 
der Kaiserlich-Koniglichen Zoologische-Botan- 
ischen Gesellschaft in Wien 47: 435-454. 

. 1909. Naucoridae. Descriptions d'especes 

nouvelles. Bulletin de la Societe des Sciences de 
Bucarest-Roumanie 18(1): 43-61. 

. 1910. Trois especes nouvelles de la famille 

Naucoridae. Bulletin de la Societe des Sciences 
de Bucarest-Roumanie 19(3): 438-444. 

Nieser, N., O. R. Gonzalez, and K. Eichelkraut. 1993. 
Nuevas especies de Naucoridae Fallen (Heterop- 
tera: Nepomorpha). Boletin del Museo Entomo- 
logia, Universidad del Valle 1: 1-11. 

Nieser, N. and P. P. Chen. 1996. Six new taxa of Ne- 
pomorpha from Sulawesi and Mindanao. Notes on 
Malesian aquatic and semiaquatic bugs (Heterop- 
tera), VI. Tijdschrift voor Entomologie 139: 161- 
174. 

Palisot de Beauvois. A. M. F. J. 1820. Insectes recueil- 
lis en Afrique et en Amerique, dans les royaumes 
dOware et de Benin, a Saint-Domingue et dans 
les Etats-Unis, pendant les annees 1786-1797. 
Fain. Paris 14: 225-240. 

Polhemus, D. A. 1994. A new species of Aphelochei- 
rus from Sumatra, and addenda to the world 
checklist (Heteroptera: Naucoridae). Journal of the 
New York Entomological Society 102: 74-78. 

Polhemus, J. T and D. A. Polhemus. 1981. Three new 
species of Amhrysus from Mexico (Hemiptera: 
Naucoridae). Pan-Pacific Entomologist 57: 397- 
401. 

. 1982. Notes on neotropical Naucoridae II. A 

new species of Amhi-ysus and review of the genus 
Potamocoris (Hemiptera). Pan-Pacific Entomolo- 
gist 58: 326-329 (1983). 

. 1988. The Aphelocheirinae of tropical Asia 

(Heteroptera: Naucoridae). Raffles Bulletin of Zo- 
ology 36: 167-300. 

. 1994. A new species of Amhjysus Stal from 

Ash Meadows, Nevada (Heteroptera: Naucoridae). 
Journal of the New York Entomological Society 
102: 261-265. 

Rawat, B. L. 1939. On the habits, metamorphosis and 
reproductive organs of Naucoris cimicoides L. 
(Hemiptera-Heteroptera). Transactions of the Roy- 
al Entomological Society of London 88: 1 19-138. 

Sites, R. W. 1990. Naucorid records from Amazonian 
Ecuador (Heteroptera: Naucoridae). Florida En- 
tomologist 73: 334-335. 

. 1991. Egg ultrastructure and descriptions of 

nymphs of Pelocoris poeyi (Guerin Meneville) 
(Hemiptera: Naucoridae). Journal of the New 
York Entomological Society 99: 622-629. 

Sites, R. W. and B. J. Nichols. 1990. Life history and 
descriptions of immature stages of Ambiysus liin- 
aiiis liiiuitns (Hemiptera: Naucoridae). Annals of 



VOLUME 101, NUMBER 



the Entomological Society of America 83: 800- 
808. 
. 1993. Voltinism, egg structure, and descrip- 



ciesque novas descripsit (continuatio). Stettin En- 
tomologische Zeitung 23: 437-462. 
. 1 876. Enumeratio Naucoridarum //; Enumer- 



tions of immature stages of Cnphocricos hunger- 
fordi (Hemiptera: Naucoridae). Annals of the En- 
tomological Society of America 86: 80-90. 

Sites, R. W. and M. R. Willig. 1994. Efficacy of men- 
sural characters in discriminating among species 
of Naucoridae (Insecta: Hemiptera): multivariate 
approaches and ontogenetic perspectives. Annals 
of the Entomological Society of America 87: 803- 
814. 

Sites, R. W., B. J. Nichols, and S. Permkam. 1997. 
The Naucoridae (Heteroptera) of southern Thai- 
land. Pan-Pacific Entomologist 73: 127-134. 

Stal, C. 1858. Bidrag till Rio Janeiro-traktens Hemip- 
ter-fauna. Konglica Svenska Vetenskaps-Akade- 
miens Handlingar 2(7): 1-84. 

. 1861. Nova methodus familias quasdam Hem- 

ipterorum disponendi. Ofversight af Kongliga 
Svenska Vetenskaps-Akademiens Forhandlingar 
18: 195-212. 

. 1862. Hemiptera mexicana enumeravit spe- 



atio Hemipterorum. Konglica Svenska Veten- 
skaps-Akademiens Handlingar 14(4): 141-147. 

Stys, P. and A. Jansson. 1988. Check-list of recent fam- 
ily-group and genus-group names of Nepomorpha 
(Heteroptera) of the world. Acta Entomologica 
Fennica 50: 1-44. 

Torre Bueno, J. R. de la. 1903. Brief notes toward the 
life history of Pelocoris femorata Pal. B. with a 
few remarks on habits. Journal of the New York 
Entomological Society 11: 166-173. 

Usinger, R. L. 1946. Notes and descriptions oi Ainb)-y- 
sus Stal with an account of the life history oi Ain- 
hrysiis mormon Montd. (Hemiptera. Naucoridae). 
The University of Kansas Science Bulletin 31: 
185-210. 

Westwood, J. O. 1833. On the connecting links be- 
tween the Geocorisae and Hydrocorisae of Latreil- 
le, or the land and water bug tribes. Magazine of 
Natural History and Journal of Zoology. Botany, 
Minerology, Geology and Meterology 6: 228-229. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999, pp. 26-38 

NEW SPECIES OF HYDROPTILIDAE (TRICHOPTERA) FROM THE 
AMAZON REGION OF NORTHEASTERN PERU 

S. C. Harris and L. J. Davenport 

(SCH) Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. (e-mail: 
harris@mail.clarion.edu); (LJD) Department of Biology, Samford University, Birming- 
ham, AL 35229, U.S.A. 



Abstract. — Two new species of Neotrichia, N. orejona and A', tirabuzona, five new species 
of Oxyethira, O. presilla, O. peruviana, O. vaina, O. picita, and O. hozosa, and one new 
species of Orthotrichia, O. shimigaya are described from the upper Amazon region in Peru. 

Key Words: Microcaddisflies, Trichoptera, Hydroptilidae, Peru, Neotropics, new species 



In 1992, Harris and Davenport described 
five new species of microcaddisflies from 
the Rio Sucusari and Rio Yanamono in the 
upper Amazon region of Peru (Fig. 1 in that 
paper). The collections were made in 1991 
by Davenport during educational expedi- 
tions to the Explorama Lodge and the Ex- 
plomapo Camp. Davenport made additional 
trips to the same region (as described in Har- 
ris and Davenport 1992) in January of 1993 
and 1995. This paper reports on seven new 
species, two in the genus Neotrichia, one in 
the genus Orthotrichia, and four in the ge- 
nus Oxyethira, from the most recent collec- 
tions. An additional Oxyethira is described 
from an earlier collection in northern Peru. 

Morphological terminology follows that 
of Marshall (1979). Length is measured 
from the top of the head to the wing tip and 
is given as a range with more than one 
specimen. Type material is deposited in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(NMNH), and in the collection of the senior 
author (SCH). 

Neotrichia orejona Harris and 
Davenport, new species 

(Fig. 1) 
Diagnosis. — In many respects, this species 
is similar to A^. yanoinonoa Harris and Dav- 



enport, A^. cayada Harris and A^. browni Har- 
ris. All three share the elongate posterolateral 
extensions from the ninth abdominal segment 
and all have a hooklike phalhc apex. The new 
species is readily separated by the triangular 
inferior appendages, the bifid subgenital plate 
and the serrated phalhc processes. 

Male. — Length 1.3 mm. Antenna with 18 
segments. Brown in alcohol. Abdominal seg- 
ment VIII annular. Segment IX in lateral view 
with narrow, elongate process from postero- 
lateral margin; in ventral view with elongate, 
narrow lobes laterally, mesally with pair of 
thin, elongate processes each bearing small 
seta at apex. Segment X fused with IX, in 
dorsal view deeply incised mesally, pair of 
small setiferous lobes anteriorly. Inferior ap- 
pendages triangular in lateral view, apex trun- 
cate bearing numerous setae; in ventral view 
rectanguloid, curving mesad. Subgenital plate 
beaklike in lateral view with narrow seta- 
bearing process dorsally; wide basally in ven- 
tral aspect with bifid processes, narrowing 
posteriorly. Phallus tubular, apex divided into 
two flattened serrate processes, ejaculatory 
duct protruding subapically, thin paramere 
encircling shelf near midlength. 

Female. — Unknown. 

Type material. — Holotype, 6 . Peru, Lor- 



VOLUME 101. NUMBER 1 



27 




Fig. 1. Neotrichia orejona. male genitalia. A. Lateral. B. Ventral. C, Dorsal. D. Phallus, dorsal. 



eto, edge of Rio Sucusari backwater, ad- 
joining Explornapo Camp, 16 January 
1993, L. J. Davenport (NMNH). 

Etymology. — Named for the Orejone In- 
dians which live in the area. 



Neotrichia tirabuzona Harris and 
Davenport, new species 

(Fig. 2) 
Diagnosis. — This new species falls with- 
in the caxima group, as established by Mar- 



28 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fis. 2. Neotrichio tirahiizomi. male genitalia. A. Lateral. B, Ventral. C. Dorsal. D. Phallus, dorsal. 



shall (1979), with greatest similarity to N. 
rotundata Flint and A'', dientera Harris. The 
hooklike inferior appendages and spiral 
process from the phallus apex are charac- 
teristic for N. tirabuzona. 

Male. — Length 1.5-1.7 mm. Antenna 



with 18 segments. Brown in alcohol. Ab- 
dominal segment VIII annular. Segment IX 
in lateral view square, curving ventrally to 
posteroventral projection; in ventral view 
square, emarginate on posterior and anterior 
margins. Segment X lobate in lateral view; 



VOLUME 101. NUMBER 1 



29 



in dorsal view partially fused with IX at 
narrow base, widening distally with trun- 
cate incision apically. Inferior appendages 
hook-shaped in lateral view; in ventral view 
square and widely separated, sclerotized 
bands basally, posteriorly, and mesally. 
Subgenital plate a narrow shelf in lateral 
view; in ventral view rounded distally, pair 
of elongate setae mesally. Phallus wide at 
base and at rounded apex, spiral process 
apically, ejaculatory duct thin and elongate, 
protruding apically, elongate paramere en- 
circling shaft beyond midlength. 

Female . — Unknown . 

Type material. — Holotype, 6 . Peru, Lor- 
eto, edge of Rio Sucusari backwater, ad- 
joining Explornapo Camp, 16 January 
1993, L. J. Davenport (NMNH). Paratypes, 
Peru, same data as holotype, 7 6 (NMNH, 
SCH). 

Etymology. — Spanish, corkscrew, refer- 
ring to the spiral process from the phallus 
apex. 

Orthotrichia shimigaya Harris and 
Davenport, new species 

(Fig. 3) 

Diagnosis. — The genus Orthotrichia is 
represented by six species in the Nearctic 
region, with two, O. aegerfasciella (Cham- 
bers) and O. cristata (Morton), extending 
into the Neotropical region. Orthotrichia 
shimigaya is the first species of the genus 
to be reported exclusively from South 
America. The species is easily recognized 
by the structure of the inferior appendages. 

Male. — Length 2.5-2.6 mm. Antenna 
with 32 segments. Brown in alcohol. Ab- 
dominal segment VII with elongate poster- 
omesal process from venter. Segment VIII 
annular. Segment IX reduced ventrally to 
narrow bridge, a rounded lobe posteroven- 
trally; in dorsal view incised posterolater- 
ally, mesally a truncate lobe. Segment X in 
lateral view divided into pair of thin, elon- 
gate processes, turned laterad at apex; in 
ventral view, narrowly incised on postero- 
lateral margin, left lobe wider and more 
truncate than right. Inferior appendages in 



lateral view short and triangular; in ventral 
view asymmetrical, left appendage with 
pair of thin, seta-bearing processes from 
posterolateral margin, right appendage nar- 
rowing mesad and curving downward, sin- 
gle seta-bearing process from posterolateral 
margin. Subgenital plate in lateral view a 
narrow shelf; in ventral view tonguelike, 
pair of setae from rounded, posterior mar- 
gin. Phallus tubular, apical half with ring- 
like crenulations, tipped with pair of narrow 
lateral lobes, paramere encircling shaft at 
midlength. 

Female. — Unknown. 

Type material. — Holotype, 6 . Peru, Lor- 
eto, small stream just outside grounds of 
Explorama Inn, 20 January 1995, L. J. Dav- 
enport (NMNH). Paratype. Peru, Loreto, 
backwater creek at outlet of Lake Shimigay, 
ca. 2 km. upstream Rio Napo from mouth 
of Rio Sucusari, 15 January 1993, L. J. 
Davenport, 1 6 (NMNH). 

Etymology. — Named for Lake Shimigay, 
one of the type localities for the species. 

Oxyethira presilla Harris and 
Davenport, new species 

(Fig. 4) 

Diagnosis. — This new species is most 
similar to O. rareza Holzenthal and Harris, 
with which it shares the asymmetrical gen- 
italic features. The multiple processes at the 
apex of the phallus and the long looping 
processes from the venter of segment IX 
will readily identify O. presilla. 

Male. — Length 2.2 mm. Antenna with 38 
segments. Brown in alcohol. Abdominal 
segment VII annular, lacking posteromesal 
process from the venter. Segment VIII ta- 
pering posteriorly, deeply incised on pos- 
terior margin in ventral and dorsal views. 
Segment IX complex, elongate anteriorly 
and tapering posteriorly with several asym- 
metrical processes, ventrally divided into 
two processes, lowermost upturned at mid- 
length and slightly widening, uppermost 
process thin, posteriorly curving into a 
loop; dorsal process narrow anteriorly, wid- 
ening to transverse plate at midlength; in 



30 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Orthotrichia shunisa.a. male genitalia. A. Lateral. B. Ventral. C. Dorsal. D. Phallus, dorsal. 



ventral view deeply incised, right lateral 
margin with two processes, apical process 
narrow, subapical process wider and pro- 
jecting more mesad, left lateral margin di- 
vided into two elongate thin processes. Ter- 



gum X apparently fused with IX as apex of 
rectangular plate on IX. Inferior append- 
ages and subgenital plate not evident. Phal- 
lus wide at base, narrowing at midlength, 
apical portion widening with several scler- 



VOLUME 101. NUMBER 1 



31 




Fig. 4. Oxyethira presillo. male genitalia. A, Lateral, left side. B, Lateral, right side. C. Ventral. D. Dorsal. 
E. Phallus, dorsal. F. Phallus, lateral. 



32 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 5. Oxyelhira peruviana, male genitalia. A, Lateral. B, Ventral. C. Dorsal. D, Phallus, dorsal. 



otized processes on lateral margins, ejacu- Type material. — Holotype, S. Peru, Lor- 

latory duct protruding distally as sclero- eto, Yanamono Creek at jungle's edge, near 

tized, sinuate process. Explorama Lodge, 12 January 1995, L. J. 

Female. — Unknown. Davenport (NMNH). 



VOLUME 101. NUMBER 1 



33 



Etymology. — Spanish, loop, refering to Etymology. — Named for the country of 

the distinctive process from segment IX. Peru. 



Oxyethira peruviana Harris and 
Davenport, new species 

(Fig. 5) 

Diagnosis. — This species appears to be 
most similar to O. spissa Kelley, a mem- 
ber of the pallida group of Kelley (1984). 
Both species have a prominent posterolat- 
eral process from segment IX and incon- 
spicuous subgenital plate, but O. peruvi- 
ana differs in the elongate processes from 
the phallus apex and the longer inferior 
appendages. 

Male. — Length 2.3 mm. Antenna with 
29 segments. Brown in alcohol. Abdomi- 
nal segment VII annular with short pos- 
teromesal process from the venter. Seg- 
ment VIII annular in lateral view; deeply 
incised posteriorly in ventral view; round- 
ed posteriorly in dorsal view. Segment IX 
triangular in lateral view, greatly reduced 
dorsally, acute, downturned curved pro- 
cess posterolaterally, posteroventrally 
with short, triangular process; in ventral 
view, posterior margins with three pro- 
cesses, lateralmost thin and acute, ventro- 
mesal processes triangular, dorsomesal 
processes tapering and angled inward. 
Segment X a short, membranous lobe; in 
dorsal view wide and rounded posteriorly. 
Inferior appendages short and rounded in 
lateral view, short spine from venter; in 
ventral view fused mesally, triangular lat- 
erally with small bilobed sclerite from 
mesal margins. Subgenital plate rectan- 
guloid and slanted posteroventrally in lat- 
eral view; in ventral view a narrow, trans- 
verse band; lacking bilobed process. Phal- 
lus tubular, widening at apex which bears 
an elongate paramere subapically and 
short, transverse process at apex; ejacu- 
latory duct bifid at apex. 

Female. — Unknown. 

Type material. — Holotype, S . Peru, Lor- 
eto, tributary to Rio Yanamono at Explor- 
ama Lodge, 11 March 1991, L. J. Daven- 
port (NMNH). 



Oxyethira vaina Harris and Davenport, 
new species 

(Fig. 6) 

Diagnosis. — This new species is most 
similar to O. orellanai Harris and Dav- 
enport, a member of the Tanytrichia sub- 
genus of Kelley (1984), which was also 
collected at the same locality on the Rio 
Sucusari. The new species is separated by 
the shorter anteroventral extension of seg- 
ment IX and the triangular inferior ap- 
pendages, which in ventral view are fused 
mesally. 

Male. — Length 2.3 mm. Antenna with 
28 segments. Brown in alcohol. Abdomi- 
nal segment VII annular with short pos- 
teromesal process from the venter. Seg- 
ment VIII tapering posteroventrally in lat- 
eral view; in ventral view deeply incised 
mesally on posterior margin; posterior 
margin of dorsum with mesal truncate in- 
cision. Segment IX reduced posterodor- 
sally to narrow bridge, narrowing and ex- 
tending anteriorly through segment VII. 
Tergum X lobate, membranous. Inferior 
appendages short and triangular in lateral 
view; fused in ventral view, round mesal 
incision creating thin triangles laterally, 
stout seta from apex, dorsal lobes along 
mesal margins. Subgenital plate in lateral 
view a narrow shelf with bilobed process 
curving over it posteriorly; in ventral 
view a narrow rectangle with thin lateral 
arms angled mesad, bilobed processes 
widely separated with setae at tips. Phal- 
lus tubular with lateral sheath, narrow 
sclerite running contiguous with ejacula- 
tory duct. 

Female. — Unknown. 

Type material. — Holotype, S . Peru, Lor- 
eto, edge of Rio Sucusari backwater, ad- 
joining Explornapo Camp, 16 January 
1993, L. J. Davenport (NMNH). 

Etymology. — Spanish, sheath, referring 
to the lateral ribbonlike band of the phallus. 



34 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Fig. 6. 




0.yer,ura raina. male genitalia. A. Lateral. B, Ventral. C. Dorsal. D. Phallus, lateral. 



VOLUME 101, NUMBER 1 



35 



Oxyethira picita Harris and Davenport, 
new species 

(Fig. 7) 

Diagnosis. — Although this species ap- 
pears to be a member of the Tanytrichia 
subgenus of Kelley (1984), there is some 
resemblance to O. zilaba (Mosely) of the 
subgenus Loxotrichia. The new species is 
distinguished by the elongate subgenital 
plate and the pair of small lateral spines on 
the tenth tergite. 

Male. — Length 2.3 mm. Antenna with 26 
segments. Brown in alcohol. Abdominal 
segment VII annular with short postero- 
mesal process from the venter. Segment 
VIII tapering posteroventrally to rounded 
apex, short process dorsolaterally; in ven- 
tral view deeply incised posteriorly; in dor- 
sal view also deeply incised with three 
acute processes mesally. Segment IX ex- 
tending anteriorly into segment VI, poste- 
riorly short and reduced dorsally to thin 
process. Segment X lobate in lateral view; 
in dorsal view rectanguloid with rounded 
apex, pair of short spines posterolaterally. 
Inferior appendages in lateral view elongate 
and tapering posteriorly, apex with ventral 
spine and dorsal seta-bearing process; in 
ventral view fused posteriorly, apex with 
lateral seta-bearing processes and median 
truncate process, heavy seta between pro- 
cesses, lateral margins gently emarginate. 
Subgenital plate curving anteroventrally, 
posteriorly elongate and narrowing to acute 
apex, transverse bilobed process thin and 
elongate; in dorsal and ventral views divid- 
ed at base into two elongate processes 
which narrow and cross apically, bilobed 
process thin, diverging distally. Phallus tu- 
bular, small lateral spines below midlength, 
widening apically, pair of lateral processes 
which are curved distally, ejaculatory duct 
between the two processes. 

Female. — Unknown. 

Type material. — Holotype, <S . Peru, Lor- 
eto, edge of Rio Sucusari backwater, ad- 
joining Explornapo Camp, 16 January 
1993, L. J. Davenport (NMNH). 



Etymology. — Spanish, small shai-p point, 
referring to the small spines of the tenth 
tergite. 

Oxyethira hozosa Harris and Davenport, 
new species 

(Fig. 8) 

Diagnosis. — This species is closely relat- 
ed to O. scaeodactyla Kelley, particularly 
in the structure of the phallus and the in- 
ferior appendages. The new species differs 
in the acute, distal point of the subgenital 
plate, the short bilobed process, and the 
more complete ninth segment. 

Male. — Length 2.4 mm. Both antenna 
broken. Brown in alcohol. Abdominal seg- 
ment VII annular with short, posteromesal 
process from the venter. Segment VIII 
rounded posteriorly in lateral view; in ven- 
tral view, deep mesal incision; emarginate 
dorsally. Segment IX tapering anteriorly, a 
narrow band posterodorsally, narrow pro- 
cess posteroventrally; ventrally with lateral 
margins produced into sharp points which 
extend beyond VIII, pair of fingerlike pro- 
cesses mesally; dorsally fused posteriorly 
with X. Segment X a short membranous 
lobe in lateral view; dorsally square in 
shape with posterior margin truncate. Infe- 
rior appendages short and truncate; in ven- 
tral view, incised mesally and fused, nar- 
rowing posterolaterally and bearing stout 
seta. Subgenital plate in lateral view strong- 
ly curving ventrad, apex with acute apical 
point; in ventral view wide with lateral edg- 
es rounded, mesally with knoblike process; 
bilobed processes short in lateral view; in 
ventral view widely separated and sinuate. 
Phallus short, distally narrowing to con- 
spicuous hook, ejaculatory duct sinuate and 
enclosed within membranous lobe. 

Female. Unknown. 

Type material. — Holotype, S . Peru, Lor- 
eto, Rio Yanamono just below Explorama 
Lodge, 10 January 1993, L. J. Davenport 
(NMNH). 

Etymology. — Spanish, sicklelike, refer- 
ring to the distinctive phallus. 



36 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. 



Oxxethira picita. male genitalia. A. Lateral. B. Ventral. C, Dorsal. D, Phallus, dorsal. 



VOLUME 101. NUMBER 1 



37 




Fig. 8. Oxyethira hozosa, male genitalia. A, Lateral. B. Ventral. C, Dorsal. D, Phallus, dorsal. 



Acknowledgments 

Thanks to the staff at Explorama Tours 
in Iquitos for their help in organizing the 
trips to Peru; to the two manuscript review- 
ers; and to Sarah Harris who typed most of 
the original draft of the manuscript. 



Literature Cited 

Harris, S. C. and L. J. Davenport. 1992. New species 
of microcaddisflies from the Amazon region, with 
especial reference to northeastern Peru (Trichop- 
tera: Hydroptilidae). Proceedings of the Entomo- 
logical Society of Washington 94: 454-470. 

Kelley, R.W. 1984. Phylogeny. morphology and clas- 



38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

sification of the micro — caddisfly genus C>.vv£'//;7ra Marshall, J. E. 1979. A review of the genera of the 

Eaton (Trichoptera: Hydroptilidae). Transactions Hydroptilidae (Trichoptera). Bulletin of the British 

of the American Entomological Society 1 10: 435- Museum (Natural History), Entomological Series 

463. 39: 135-239. 



PROC. ENTOMOL. SOC. WASH. 
101(1), 1999. pp. 39-56 

AN EXAMINATION OF THE NORTH AMERICAN APHID SPECIES 

CURRENTLY PLACED IN OVATUS VAN DER GOOT (HEMIPTERA: 

APHIDIDAE) WITH THE DESCRIPTION OF A NEW GENUS 

Andrew S. Jensen and Manya B. Stoetzel 

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department 
of Agriculture, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: ajensen@ 
sel.barc.usda.gov; mstoetze@sel.barc.usda.gov) 



Abstract. — The current placement in Ovatus of four endemic North American aphid 
species is critically examined. Results of cladistic analyses clearly indicate that these four 
species are not congeneric with Palearctic Ovatus and that they are more closely related 
to Myzus Passerini. Cladistic evidence is provided to justify the erection of a new genus, 
Abstriisomyzus n. gen., for the four endemic North American species formerly placed in 
Ovatus. This resulted in four new combinations: Abstrusomyzus leucocrini (Gillette & 
Palmer) n. comb., Abstrusomyzus phloxae (Sampson) n. comb., Abstrusomyzus reticulatus 
(Heie) n. comb., and Abstrusomyzus vahiliae (Robinson) n. comb. Information on each 
of the four species is compiled and presented. Illustrations are provided, along with a key 
to the species of the new genus and notes on the single remaining species of Ovatus in 
North America, Ovatus crataegarius (Walker). 

Key Words: polyphagy, Myzus, Ovatus, new genus, cladistics 



Most aphids are extremely host specific, 
feeding on one or a few species of plants 
that are usually closely related (Eastop 
1973). Many of the most pestiferous aphids 
known differ dramatically from this normal 
pattern by being polyphagous on plants in 
widely divergent families. Well-known po- 
lyphagous pests include the green peach 
aphid {Myzus persicae (Sulzer)], the potato 
aphid [Macrosiphum euphorbiae (Thomas)], 
and the cotton aphid {Aphis gossypii Glov- 
er). These aphids can cause serious problems 
even when they feed on a crop in low num- 
bers, since they can transmit plant viruses 
between their phylogenetically disparate 
host plants. Very few polyphagous aphids 
are not pests. Given the fact that many of 
the most serious pest aphids are polypha- 
gous, it is important to study newly-discov- 
ered cases of polyphagy in aphids. As po- 



lyphagous aphids are likely to some day be- 
come pests, prior knowledge of their biolo- 
gy, taxonomy, and evolution will facilitate 
an effective reaction to their emergence as 
pests. 

Ovatus phlo.xae (Sampson) is one of the 
few polyphagous aphids that so far is not a 
pest. It was described in 1939 from Phlo.x 
subulata from Berkeley, California. Since 
then it has been found on eighteen plants in 
fourteen families (Table 1). It has been col- 
lected in most regions of the United States 
and probably occurs in most temperate parts 
of North America. Most plants on which this 
aphid has been found have a basal rosette of 
leaves, or tend to be prostrate. The aphid 
usually lives in the crown or on the lower 
leaves. It is possible that this species is more 
habitat specific than host specific, much as 
in Rhopalosiphum uymphaeae (L.), which 



40 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Table 1. Host plants and distributions for five species heretofore placed in Ovatus. 



Aphid Species 



Host Plants 



Distribution 



Ovatus crataegarius (Walker) 



Ovatus leucocrini (Gillette & Palmer; 
Ovatus phloxae (Sampson) 



Ovatus reticulotus Heie 
Ovatus valuliae (Robinson) 



Rosaceae 


Almost 


Crataegus spp. 


worldwide. 


Cydonia spp. 


with a 


Malus spp. 


Palearctic 


Lamiaceae 


origin 


Mentha spp. 




some other Lamiaceae 




Liliaceae 


Colorado 


Leucocritium montanum 




Apocynaceae 


Canada: 


Apocynum androsaefolium 


British 


Apocynum sp. 


Columbia 


Asteraceae 




Achillea sp. 


U.S.A.: 


Agoseris sp. 


California 


Centaurea sp. 


Colorado 


Brassicaceae 


Maryland 


Capsella hursa-pastoris 


Mississippi 


Caryophyllaceae 


Nebraska 


Cerastium vulgatum 


Oregon 


Stellaria crispa 


Pennsylvania 


Cyperaceae 


Utah 


Carex densa 


Virginia 


Fabaceae 




Trifolium sp 




Hydrophyllaceae 




Phacelia nemoralis 




Liliaceae 




"Lilies" 




Plantaginaceae 




Plantago major 




Polygonaceae 




Polygonum paronychia 




Ranunculaceae 




Ranunculus sp. 




Polemoniaceae 




Phlox subulata 




Rubiaceae 




Galium sp. 




Violaceae 




Viola sp. 




Oxalidaceae 


North Carolina 


Oxalis '.'stricta 




Rosaceae 


Manitoba 


Fragaria vesca 





feeds widely on aquatic and semiaquatic 
plants. If this were true, O. phloxae could 
emerge at any time as a pest of a crop with 
an appropriate growth habit. Therefore a 
study of North American Ovatus was un- 
dertaken to provide needed information re- 



garding O. phloxae and three closely related 
endemic North American aphids currently 
placed in Ovatus. The relationship of these 
four species to Palearctic Ovatus is exam- 
ined using cladistics, and a new genus is 
proposed for them based on these analyses. 



VOLUME 101, NUMBER 1 



41 



Materials and Methods 

Most specimens studied are housed in the 
National Collection of Aphidoidea (USNM, 
located at the Systematic Entomology Lab- 
oratory, USDA, Beltsville, Maryland, USA). 
Others were obtained on loan from The Nat- 
ural History Museum, London (BMNH); 
Agriculture Canada, Vancouver, British Co- 
lumbia (UBC); University of California, 
Berkeley (UCB); and Oregon State Univer- 
sity, Corvallis (OSU). 

Aphids were mounted on microscope 
slides in Canada balsam or Hoyer's medium, 
and observed under phase contrast micros- 
copy. Terminology follows Miyazaki (1987). 
Drawings were made by the first author us- 
ing a camera lucida. 

Cladistic analyses were performed using 
PAUP 3.1.1 (Swofford 1993) and MacClade 
3.01 (Maddison and Maddison 1992) on 23 
taxa using 25 characters. Apterous and alate 
viviparae were included in the character 
analysis, supplemented by one character of 
the first instar nymph. All characters were 
treated as unordered. 

Ovatus van der Goot 

Ovatus has most recently (Remaudiere 
and Remaudiere 1997) been used for four- 
teen MycM^-like aphids, which generally mi- 
grate between Pomoidea and various Lami- 
aceae, or are monoeceous on either of these 
groups. Nine of the fourteen species fit this 
pattern of host plant association. One excep- 
tion is the single member of the subgenus 
Ovatoides, Ovatus (Ovatoides) inulae 
(Walker). This species feeds on Asteraceae 
and differs morphologically from most Ova- 
tus by the nearly complete lack of spinula- 
tion on the head and first two antennal seg- 
ments, and the long, setose ultimate rostral 
segment. The other four species that do not 
conform with the Pomoidea/Lamiaceae host 
plant association are the four endemic North 
Amencan species [Table 1, excluding Ova- 
tus crataegarius (Walker)]. These species 
feed on herbs from a diverse set of plant 
families, and none of them is known from 



Pomoidea or Lamiaceae. Morphologically 
these species differ from most Palearctic 
Ovatus by the dark bars and blotches on the 
abdominal tergum of the alate vivipara 
(Figs. 17, 18), first tarsal chaetotaxy of 3,3,2, 
and the peculiarly shaped siphunculus (Figs. 
9-12). 

The first to place one of these species in 
Ovatus was Hille Ris Lambers (1966), who 
transferred Phorodon phloxae to Ovatus 
without any explanation. Heie (1972) de- 
scribed Ovatus reticulatus, and Eastop and 
Hille Ris Lambers (1976) transferred Myzw^ 
leucocrini Gillette and Palmer and Myzus 
valuliae Robinson without explanation. The 
obviously close relationship among these 
four species dictated that they be placed to- 
gether in the same genus, but their morpho- 
logical and biological differences from Pa- 
learctic Ovatus causes doubt about their cur- 
rent generic placement. 

Cladistic Analyses 

This study attempts to determine whether 
the four North American species currently 
placed in Ovatus will cluster together, and 
whether they will form their own group, 
form a part of Ovatus, or fall within another 
genus. Doing this requires that the cladistic 
analysis include several species from one or 
more genera to which the species in question 
might be more closely related than they are 
to Ovatus. These cladistic analyses are 
meant to explore the proper generic place- 
ment of the four endemic North American 
species currently placed in Ovatus and are 
not meant to resolve species-level issues for 
the other aphids included in the analysis. 

Choice of appropriate comparative groups 
for a test of the proper placement of the four 
endemic North American species is partic- 
ularly troublesome in the case of Ovatus for 
at least three reasons. First, the fact that the 
analysis deals with aphids presents problems 
in itself (see Jensen 1997 for some discus- 
sion of this issue). Aphids are highly pro- 
genetic sensu Gould (1977), and because of 
their conservative morphology most genera 
lack apomorphic characters. Instead, there is 



42 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



essentially a large pool of characters that oc- 
cur across a wide range of aphid taxa, and 
many genera are recognized on the basis of 
a unique combination of these characters. 
Aphid genera are also frequently defined by 
the absence of one or more characters typi- 
cal in genera to which they are most similar 
morphologically. These facts, in combina- 
tion with the lack of work on the evolution 
of the commonly used aphid characters, 
make deductions about relatedness difficult. 

The second problem that Ovatus poses is 
biogeographical. As currently understood, 
the genus is Holarctic, with ten species na- 
tive to Eurasia, and four native to North 
America. Perhaps, then, comparative species 
should be drawn from related genera on both 
continents. Nearly all species of Myzina, the 
subtribe to which Ovatus belongs, are en- 
demic to Eurasia, with the subtribe's greatest 
diversity in southern and eastern Asia. 
Therefore, choosing Eurasian species for 
comparison is easy. But examination of the 
native aphid fauna of North America shows 
that there are only a handful of Myzina na- 
tive to this continent. These native species 
are mostly speciahzed moss and sedge feed- 
ers such as Myzodium Bomer and Caroli- 
naia Wilson. There are few other native 
North American Myzina besides the four pu- 
tative Ovatus, and several species of Hyalo- 
myzus. In other work (Stoetzel, Miller and 
Jensen, in preparation), evidence has been 
found for the monophyly of Hyalomyzus 
Richards of North America. This leaves few 
North American groups to which the four 
Ovatus could belong. 

Thirdly, the four North American species 
currently placed in Ovatus are extremely 
similar and possess some unique character- 
istics. For example, they all have siphunculi 
more or less cylindrical, but slightly expand- 
ed apically (Figs. 9-12). Two of the species 
also tend to have more than three pairs of 
setae on the cauda, whereas most Myzina 
have two or three pairs. The four North 
American species also have a distinctive pat- 
tern of pentagonal or hexagonal reticulation 
on their terga in the apterous viviparous fe- 



male. These peculiarities make them isolated 
morphologically from most of the Myzina 
and make the choice of appropriate species 
for the current analysis more difficult. 

Myzus has for a long time been used for 
a great diversity of species of Myzina. This 
can be illustrated by examining the list of 
Eastop and Hille Ris Lambers (1976). They 
list 53 valid species of Myzus, and 40 others 
that were described in Myzus, but subse- 
quently transferred to 27 other genera. The 
range of species included in the genus has 
varied from one author to another during this 
century. In North America most workers 
have dealt with the genus in a broad sense 
(Mason 1940, Palmer 1952). Mason states 
that Myzus can be recognized among the 
Macrosiphina in his tribe Aphidini by the 
convergent frontal tubercles. Thus in his re- 
vision of the genus. Mason (1940) included 
20 species that are today scattered among six 
genera. Despite all this modification or re- 
striction of the definition of the genus, My- 
zus still contains a wide array of species and 
is quite likely polyphyletic. The diversity of 
species in Myzus and the lack of another ge- 
nus that is cleaiiy related to the four species 
in question indicate that Myzus is the best 
choice for use in this analysis. Myzus pro- 
vides a wide array of species to which the 
four endemic North American species might 
prove to be closely related. 

Species Included in the Analyses 

Included in the analysis were Ovatus 
phloxae and three very similar North Amer- 
ican species, O. leucocrini, O. reticulatus, 
and O. valuliae. For purposes of testing their 
relationship to Palearctic Ovatus, six other 
Ovatus species were studied, including Ova- 
tus (Ovatoides) inulae, the only member of 
its subgenus. These were chosen based on 
their availability in the collection of the 
USNM and material obtained on loan. Thus 
ten of the world's fourteen Ovatus species 
were analyzed. Adequate material of the oth- 
er Eurasian species currently placed in Ova- 
tus was not available. 

Ten species of Myzus were included. 



VOLUME 101, NUMBER 1 



43 



Table 2. Data matrix used in the cladistic analyses. Question marks indicate missing data, t indicates a 0/1 
polymorphism, and a t indicates a 0/2 polymorphism. 



111111111 122222 2 
123456789012345678 901234 5 



H. eriohotrycw 

H. jiissiaecie 

H. luitchellensis 

M. ascalonicus 

M. cerasi 

M. certiis 

M. cymhalariae 

M. ciycei 

M. hemerocallis 

M. lytliii 

M. ornatiis 

M. persicae 

M. varians 

O. crataegarius 

O. glechomae 

O. insitus 

O. inulae 

O. leucocrini 

O. malisuctits 

O. menthahiis 

O. plilo.xoe 

O. reticulatus 

O. valitliae 



00000 
00000 
00011 
t201t 
00000 
00011 
12010 

oooot 

02011 
00001 
10011 
00010 
00000 
00110 
10010 
00110 
OHIO 
03110 
OHIO 

tooio 

00010 
03010 

13010 



00000 
00001 
10100 
01010 
00000 

oono 

00011 
00011 

ootio 

00000 
00000 
00210 
00010 
00001 
00010 
00011 
11010 
01010 
00010 
00000 
00011 
00210 
00010 



00001110 
00001110 
00101110 
10101110 
00000000 
10101010 
10001110 
10100110 
00100010 
00100010 
10110110 
10001010 
10100110 
00000110 
00010110 
00000110 
00110010 
11102001 
lOlOOltO 
OOltOllO 
1110201t 
11102000 
111120C0 



111011 

110011 ? 

111011 

tttOOO 1 

000000 

000100 1 

010000 1 

010110 1 

000011 

000000 

000110 

000100 1 

000110 1 

111101 1 

110001 1 

111101 1 

111101 1 

000100 1 

011110 1 

111001 1 

OtOOOt 1 

? ?? ?0 ? 1 

000100 1 



representing three of four subgenera, and 
16% of the world fauna. These species 
were chosen to represent all possible sub- 
genera, and are all the species of which 
adequate material was available. Three 
outgroup species were chosen from North 
American Hyalomyzus. The dataset is 
shown in Table 2. 

Characters 
Apterous viviparae 

Head capsule: 

1. Anterior setae on dorsum of head cap- 
sule: less than 0.5 times basal width of 
antennal segment III (0); more than 0.5 
times basal width of antennal segment 
III (1). 

2. Ornamentation on dorsum of head cap- 
sule: spinules or nodules present anteri- 
orly and along margins, smooth in mid- 
dle posteriorly (0); without spinules or 
nodules, sometimes with wrinkles (1); 
spinules or nodules entirely covering 



dorsum of head (2); dorsum of head with 
posterior surface in middle with polyg- 
onal reticulation (3) (Figs. 5-8). 

3. Venter of head capsule adjacent to eyes: 
spinulose (0); smooth (1). 

4. Ventral tubercles at rear of head capsule: 
present (0); absent (1). 

Antenna: 

5. Ratio of processus terminalis to the base 
of antennal segment VI: greater than 3 
(0); less than 3 (1). 

6. Venter of antennal segment II: bumpy 
(0); smooth (1). 

Mouth Parts: 

7. Pairs of setae on rostral segment III: al- 
most always 2 (0); more than 2(1). 

8. Ultimate rostral segment: longer than 
hind tarsal segment II (0); shorter than 
hind tarsal segment II and without ac- 
cessory setae ( 1 ); shorter than hind tarsal 
segment II and with accessory setae (2). 

9. Number of setae on ultimate rostral seg- 



44 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



{77} 
53 



{52} 
62 



{2} 



(2) 



{54} 



I M ascalonicus 

{77} M. cymbalariae 

^^ ' M. cerasi 



OZ I 



(85) 

77 



{2! 



M. lythh 
M. hemerocallis 
M. certus 
M. persicae 
O. leucocrini 
O. reticulatus 
O. valuliae 
O. phloxae 
M. ornatus 
M. varians 
M. dycei 
O. malisuctus 
O. glechomae 
O. crataegarius 
O. insitus 
O. inulae 
O. mentharius 
H. mitchellensis 
■ H. jussiaeae 
H. eriobotryae 

Fig. L Tree number one of two equally parsimonious trees found using the heuristic search option in PAUP 
with successively weighted data (weighted consistency index: 0.67; retention index: 0.84; and rescaled consis- 
tency index: 0.57). This tree was also one of 135 trees found using unweighted data (unweighted consistency 
index: 0.47; retention index: 0.64; and rescaled consistency index: 0.30). Numbers below some branches indicate 
branch support values from decay analyses, plain numbers indicating unweighted branch support, and numbers 
in braces indicating rescaled branch support from the weighted data (Bremer 1994). Numbers above branches 
display results of bootstrap analyses, plain numbers indicating results using unweighted data, and numbers in 
braces representing weighted data. 




ment: 0-3 (usually 2) or rarely 4 acces- 
sory setae (0); usually 3 or more (rarely 
2) accessory setae (1). 

Legs: 

10. Dorsal base of hind tibia: smooth or 
slightly wrinkled (0); distinctly sca- 
brous or imbricated (1). 

1 1 . First tarsal segments chaetotaxy for- 
mula: 3-3-3 (0); 3-3-2 (1). 

Abdomen: 

12. Tergum patterning or sculpturing: 
maze-like with no distinct polygons, or 



irregular closed shapes with smooth 
space between them (0) (Fig. 3); 
smooth (1); pentagonal or hexagonal 
(2) (Fig. 4). 

13. Spinal tubercles on abdominal tergite 
VII: usually present (0); absent (1). 

14. Setae on abdominal tergite VIII: much 
shorter than basal width of antennal 
segment III (0); subequal to basal width 
of antennal segment III (1). 

15. Shape of siphunculi: cylindrical or ta- 
pering (0); distinctly swollen (1); very 
slightly expanded near apex (2). 



VOLUME 101. NUMBER 1 



45 



O. leucocrini 
O. reticulatus 
O. valuliae 
O. phloxae 
M. certus 
M. persicae 
M. ascalonicus 
M. cymbalariae 
M. cerasi 
M. lythri 
M. hemerocallis 
M. ornatus 
M. varians 
M. dycei 
O. malisuctus 
O. glechomae 
O. crataegarius 

Fig. 2. Tree number one of four equally parsimonious trees found using the heuristic search option in PAUP 
with successively weighted data (weighted consistency index: 0.77; retention index: 0.84; and rescaled consis- 
tency index: 0.65). This tree was one of 14 trees found using unweighted data {unweighted consistency index: 
0.47; retention index: 0.64; and rescaled consistency index: 0.30). Numbers below some branches indicate branch 
support values from decay analyses, plain numbers indicating unweighted branch support, and numbers in braces 
indicating rescaled branch support from the weighted data (Bremer 1994). Numbers above branches display 
results of bootstrap analyses, plain numbers indicating results using unweighted data, and numbers in braces 
representing weighted data. 





{53) 

r+- 

{2} 


f831 


(89) 

76 
1 






H- 
{1} 


52 

r-H 
1 


1 






{6} 












-1- 
(2) 


{77) 

52 

1 






{93} 

51 

[4- 

(6) 


H- 

(2) 


(2 

{67) 
(4) 


} 

{65) 

-1- 
(2) 


{73} 

65 

(H- 

11) 








1 






1 
1 




(1) 

















16. Apical spinulation of cauda: entire, 
without blank spaces dorsally (0); re- 
duced with blank spaces dorsally (1). 

17. Abdominal tergum pigmentation: pig- 
mented, with complete or nearly com- 
plete dorsal shield (0); pale, or with 
only cross bands on tergites VII and/or 
VIII. 

18. Setae on cauda: 2 or 3 lateral pairs (0); 
4 or more lateral pairs (1). 

Alate viviparae 



Antenna: 

19. Secondary rhinaria on antennal 
ment V: absent (0); present (1). 

20. Secondary rhinaria on antennal 
ment IV: absent (0); present (1). 

21. Secondary rhinaria on antennal 
ment III: restricted to approximately 
half the circumference of the segment 



seg- 



seg- 



seg- 



(0); distributed around the entire cir- 
cumference of the segment (1). 

Mouthparts: 

22. Ornamentation of lateral part of man- 
dible (base of mouthparts): spinulose or 
scabrous (0); smooth (1). 

Abdomen: 

23. Lateral abdominal tubercles: present in 
at least some specimens (0); absent (1). 

24. Pigmented abdominal patch or bands: 
present (0); absent (1). 

Nymphs 

Legs: 

25. Apical spinulation of hind tibia: present 
(0); absent (1). 

Cladistics Results and Discussion 

The full data set was analyzed using the 
heuristic search option in PAUP. One hun- 



46 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 3. 4. Dorsal sculpturing. 3, Ovatus crataegahus. 4, Abstnisumyzus leucochni. 



dred iterations of random addition sequence 
were performed, finding 135 equally parsi- 
monious trees of length 91 steps. These had 
a consistency index (CI) of 0.47, retention 
index (RI) of 0.64, and rescaled consistency 
index (RC) of 0.30. One of these trees is 
presented in Fig. 1, with numbers below a 
few branches indicating results of a decay 
analysis (see Bremer 1994), and numbers 
above branches indicating the level of boot- 
strap support from 100 replications of heu- 
ristic bootstrap searching. Only branches 
supported by more than 50% of the boot- 
strap trees are labeled in Fig. 1. In all 135 
trees, the group of four North American 
Ovatus is placed among the Myzus species, 
separate from other Ovatus, and supported 
by more than 50% of the bootstrap trees. 
Old World Ovatus are placed as close rel- 
atives to the outgroup Hyalomyzus. The 
close relationship between Ovatus and Hy- 
alomyzus has been discussed by Eastop 
(1966), Nielsson and Habeck (1971), and 
Blackman and Eastop (1994). 

The four North American Ovatus formed 
a clade in all 135 trees and were often 
placed with M. certus and M. persicae, two 



species with swollen siphunculi that repre- 
sent the subgenus Nectarosiphon. Moving 
the four North American species from with- 
in Myzus to the more basal parts of the tree 
along with the other Ovatus caused an in- 
crease in tree length of three or four steps, 
depending on the branch to which they 
were attached. For example, moving the 
branch composed of these four species to 
become the terminal Ovatus, along with 
Ovatus malisuctus (Matsumura), caused an 
increase in tree length of three steps. But it 
is clear from the bootstrap results that the 
most strongly supported clade of more than 
two species is that composed of the four 
North American species related to O. phlox- 
ae, henceforth referred to as the '^phloxae 
group." 

A successive weighting procedure using 
the default settings in PAUP was conduct- 
ed, finding two of the 135 trees found using 
unweighted data, with the following 
weighted statistics: length, 22, 140; CI, 
0.67; RI, 0.84; RC, 0.57. The tree shown in 
Fig. 1 is one of these trees. Bootstrap and 
decay analyses were performed using the 
weighted dataset, and their results are dis- 



VOLUME 101, NUMBER 1 



47 



played in Fig. 1. Decay indices in braces in 
Fig. 1 are rescaled (Bremer 1994). The two 
successive weighting trees even more 
strongly support the monophyly of the 
phloxae group and its separation from Pa- 
learctic Ovatus. Monophyly of the phloxae 
group was supported in 78% of the boot- 
strap trees and through 2 rescaled steps of 
decay. 

The results of this analysis clearly indi- 
cate that the phloxae group does not belong 
to Ovatus. Consistent placement of the 
phloxae group within the clade of Myzus 
species indicates that they are more closely 
related to Myzus than they are to Ovatus of 
the Palearctic. It should also be pointed out 
that O. malisuctus is placed among the My- 
zus clade in some of the 135 trees, and may 
be more closely related to Myzus than are 
the other Ovatus. Further studies focusing 
on Palearctic aphids in and related to Ova- 
tus will be required to finalize the classifi- 
cation of this and other problematic species 
not included in this study. 

The unique characters of the four North 
American "Ovatus" species (peculiar si- 
phuncular shape, generally more setose ros- 
trum and Cauda, reticulate tergum) may jus- 
tify the erection of a new genus. Therefore, 
another analysis was conducted focusing 
only on the relationship of these four spe- 
cies to the Myzus included in the analysis. 
The objective of this analysis was to deter- 
mine whether the phloxae group and the 
available Myzus species would form sepa- 
rate monophyletic groups. The same set of 
characters was used. Three Palearctic Ova- 
tus were included, the first two as outgroup 
taxa: O. crataegarius, Ovatus glechomae 
Hille Ris Lambers, and O. malisuctus. In- 
group species were the phloxae group, and 
all ten Myzus in the first analysis. 

A "branch and bound" search found 
fourteen trees of length 71, with a CI of 
0.55, RI of 0.56, and RC of 0.31. One of 
these trees is presented in Fig. 2. Similar to 
the previous analysis, the phloxae group is 
part of a clade with two Myzus (Nectaro- 
siphon) species. These Myzus have swollen 



siphunculi and, similar to the phloxae 
group, two setae on the first segment of the 
hind tarsus. Rearranging the trees to make 
the phloxae group and Myzus monophyletic 
requires only a two step increase in tree 
length. A successive weighting analysis 
was conducted, which selected four of the 
original fourteen trees of length 71 (weight- 
ed statistics: length, 19, 611; CI, 0.77; RI, 
0.84; RC, 0.65). The tree in Fig. 2 is one 
of these trees. The results of weighted and 
unweighted decay and bootstrap (500 rep- 
lications) analyses are presented in Fig. 2 
as well. Unweighted data supported the uni- 
ty of the phloxae group through 1 step of 
decay and in 52% of the bootstrap trees. 
Few branches were strongly supported by 
either measure using unweighted data. 
Weighted data yielded very strong support 
for the phlo.xae group, which held up 
through six steps of rescaled decay and was 
present in 83% of the bootstrap trees. 

These results point to the necessity for 
the erection of a new genus to house the 
phloxae group. First of all, it is clear that 
the phloxae group is not congeneric with 
Ovatus from the Palearctic. The full anal- 
ysis that used Hyalomyzus as outgroup was 
overall rather weak, but where it was rela- 
tively strong was in the separation between 
the basal Hyalomyzus/Ovatus clade and the 
more apical Myzus/phlo.xae group clade. 
This showed that the phloxae group is more 
closely related to Myzus than to Palearctic 
Ovatus. The next step in the analysis ex- 
amined more closely the relationship be- 
tween the phloxae group and the other My- 
zus. The results showed that there was 
strong support for the monophyly of the 
phlo.xae group, but not for the relationship 
between it and the Myzus with which it was 
placed in the trees. Thus not only is there 
no support for the placement of the phlo.xae 
group in Ovatus, but the group also appears 
to be only weakly related to any species of 
Myzus. Our cladistic results as well as the 
peculiar morphology of the phloxae group 
provide ample justification for the erection 
of a new genus, as we do below. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Abstrusomyzus Jensen and Stoetzel, 
new genus 

(Figs. 4-18) 

Type species: Phorodon phloxae Sampson 

Diagnosis. — Abstrusomyzus can be sep- 
arated from Ovatus, Myzus and other My- 
zus-like genera in North America by the pe- 
cuHar shape of the siphunculi in all stages 
(Figs. 9-12), the distinctly pentagonal/hex- 
agonal reticulated pattern of the tergum 
(Fig. 4), 3,3,2 first tarsal segment chaeto- 
taxy, and the usual presence of pigmented 
cross-bands on the abdomen of alate vivi- 
parae (Figs. 17, 18). The completely pig- 
mented tergum in apterous viviparae of the 
three species other than A. phloxae is also 
unusual. One North American species that 
may be closely related to Abstrusomyzus is 
Aphthargelia symphoricarpi (Thomas). The 
latter species shares with Abstrusomyzus the 
same type of dorsal reticulate sculpturing, 
moderately prominent Myzus-Xike antennal 
tubercles, a relatively large number of cau- 
dal setae, large lateral tubercles, and dark 
transverse abdominal bands in the alate vi- 
viparae. Abstrusomyzus and A. symphori- 
carpi differ significantly, in that the latter 
species has 3,3,3 first tarsal chaetotaxy, 
very short almost barrel-shaped siphunculi, 
and more numerous secondary rhinaria on 
antennal segments III-V. 

Description. — Apterous vivipara: Nymph: 
hind tibiae of nymphs without spinules api- 
cally. Adult: body length 0.83-1.94 mm. 
Dorsum of head capsule (Figs. 5-8) pale to 
black, with small sparse spinules often ar- 
ranged in curving rows. Spinal region of 
head capsule with more or less triangular 
area of reticulate sculpturing; spinal tuber- 
cles sometimes present. Dorsum of head 
normally with 4 setae in posterior row, with 
3 pairs farther forward; small lateral ocelli 
or traces thereof often present. Antennal tu- 
bercles moderately to strongly produced, 
rough, with several setae. Frons often pro- 
duced slightly as a median tubercle. Setae 
on head short, blunt, much shorter than bas- 
al width of antennal segment III. Eyes nor- 



mal, with ocular tubercles and interfacetal 
spaces brownish pigmented. Ventral surface 
of head capsule more or less evenly, but 
sometimes very lightly, spinulose, with spi- 
nules often arranged in rows. Antennal tu- 
bercles with large ventral projections bear- 
ing a few short setae. Ventral head setae 
sometimes as short as the dorsal setae, 
sometimes about twice as long. Mandibular 
area of mouthparts smooth. Antennae nor- 
mally shorter than body; antennal segment 

I rough, roughest medially, with several se- 
tae, these about equal in length to those on 
dorsum of head capsule; antennal segment 

II rough, roughest ventrally and medially, 
with course imbrications; antennal segment 

III covered with imbrications, rarely with 1 
or 2 secondary rhinaria; remainder of an- 
tenna roughly imbricated. Rostrum reaching 
middle of thorax, with segment II strongly 
ornamented with rows of spinules; segment 
III with 2 pairs of setae, these located in 
apical Vi; ultimate rostral segment about 
equal in length to tarsal segments II, or 
about 0.1 mm, with 3-9 accessory setae. 
Pronotum with 3 pairs of short setae: 1 spi- 
nal, 1 pleural, and 1 lateral; the lateral pair 
associated with a frequently present pair of 
lateral tubercles that are sometimes bi- or 
trifid; setae about equal in length to those 
on the dorsum of the head. Surface of pron- 
otum reticulate, the reticulations usually 
somewhat flattened front to back. Meso- 
and metathoracic terga reticulate and often 
pigmented, setae extremely short, blunt. 
Coxae of all legs covered with spinulose 
imbrications. Femora with sparse imbrica- 
tions, especially anteriorly, with setae about 
equal in length to dorsal head setae. Tibiae 
sparsely setose, basal setae much shorter 
than those at apex. First tarsal segments 
with 3,3,2 setae (i.e., hind tarsus I with 2 
setae), second tarsal segments imbricated. 
Mesothoracic furca sessile or with short 
stalk. Abdomen with tergum reticulate 
throughout, sometimes pigmented. Dorsal 
abdominal setae blunt, very short, except 
sometimes longer on tergite VIII. Abdom- 
inal segments II-VI often with lateral tu- 



VOLUME 101, NUMBER 1 



49 




111 I I I ' I II I 'I'l ' 'r *" ^ ' I I I I 



0.180 mm 




miiLiI^S^^i^^ 



0.170 mm 



j?22222nn2 



0.156 mm 




Figs. 5-8. Head with first three antennal segments of apterous viviparae. 5, Abstrusomyzns leucocriiii. 6, A. 
phloxae. 7, A. reticulatus. 8, y4. vahdiae. 



50 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 9-16. Siphunculi and caudas of apterous viviparae. 9. L^, Ahsrrusoniyziis leucocrini. 10, 14, A. phloxae. 
11, 15, /4. reticulatus. 12, 16, A. valuliae. 



bercles of various sizes. Siphunculus cylin- 4-15 setae, more or less pointed apically. 

drical over most of its length, slightly swol- Tergite VIII with 4-7 setae, normally 4. 

len toward the tip (Figs. 9-12), imbricated Ventral abdominal setae pointed. Gonapo- 

throughout. Cauda moderately long, with physes 3 in number. 



VOLUME 101, NUMBER 1 



51 










3^ r'^'^iCpO ^'^'^'^ 



0.270 



Figs. 17, 18. Abdominal teigum of alate viviparae. 17, Abstnisoniyzns leiicocrini. 18. A. phloxae. 




Alate vivipara: Body length 0.98-1.92 
mm. Head capsule dark, slightly wrinkled 
or smooth, not reticulate. Lateral and me- 
dian ocelli present, median ocellus creating 
a strong median tubercle. Antennal tuber- 
cles only slightly rough, with several setae. 
Ventral surface of head capsule more or less 
smooth, sometimes with a few sparse spi- 
nules. Antennal segment III with secondary 
rhinaria scattered over most or all of its 
length, limited to one side, normally less 
than 20 in number; antennal segment IV oc- 
casionally with a few secondary rhinaria; 
antennal segment V without secondary rhi- 
naria. Thoracic tergites smooth. Sclerites of 
thorax dark brown, of normal design. Ab- 
domen with dark lateral sclerites, and often 
dark cross bands on some segments. Dark 
abdominal cross bands often reticulate in a 
similar fashion to the tergum of apterous 
vivipara. Otherwise essentially as in apter- 
ous vivipara. 

Etymology. — The generic name is taken 
from the Latin word "abstrusus," meaning 
hidden or concealed and Myzus, which is 
based on the Greek "myzo," meaning suck. 
The name is meant to draw attention to the 



way the type species of the genus is often 
hidden in ant-created shelters or on the low- 
er leaves of its host plants. The gender is 
masculine. 

Key to Apterous Viviparae of 
Abstrusomyzus 

1. Dorsum of apterous vivipara unpigmented; an- 
tennal tubercles strong, with prominent con- 
verging processes (Fig. 6); dorsum with hex- 
agonal reticulation usually faint due to the pale, 
unsclerotized tergum 

AbsfnisoniyzKS phloxae (Sampson) 

- Dorsum of abdomen of apterous vivipara dark 
pigmented, usually black: antennal tubercles 
strong, but without converging processes (Fig. 
5, 7. 8); dorsum with distinct hexagonal retic- 
ulation (Fig. 4) 2 

2. Apterous vivipara without traces of lateral 
ocelli, and without spinal tubercles on head; 
antennal segments I & II nearly black, antennal 
segment III unpigmented, except extreme tip 
slightly dusky, dorsum entirely dark, concolo- 
rous with cauda and siphunculus 

Abstrusomyzus reticulatus (Heie) 

- Apterous vivipara usually with traces of lateral 
ocelli, and sometimes with low spinal tubercles 
on head; antennal segment I brown, antennal 
segment II much paler than I, antennal segment 
III with tip brown; dorsum entirely dark or 



52 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



with some pale areas, cauda and siphunculus 
pigmentation variable 3 

3. Tergite VIII with middle pair of setae pointed, 
about twice as long as lateral pair of setae on 
tergite VIII, and about three times as long as 
setae on tergite VII; cauda usually with fewer 

than 9 setae (Fig. 16) 

Abstrusomyzits valulicte (Robinson) 

- Tergite VIII with middle pair of setae blunt, 
about equal in length to the lateral pair on ter- 
gite VIII and those on tergite VII; cauda usu- 
ally with more than 9 setae (Fig. 13) 

. . Ahslrusomvzus leiicocriiii (Gillette and Palmer) 



Key TO Known Alate Viviparae of 
Abstrusomyzus 

1. Setae on abdominal tergite VIII distinctly lon- 
ger than setae on tergite VII, and about equal 
to basal width of antennal segment III; lateral 
tubercles small or absent 

Abstrusomyzus valuliae (Robinson) 

- Setae on abdominal tergite VIII about equal in 
length to the setae on abdominal tergite VII, 
and much shorter than the basal width of an- 
tennal segment III; lateral tubercles various . . 2 

2. Cauda relatively setose, usually with more than 
9 setae; lateral abdominal tubercles relatively 
large (Fig. 17); antennal tubercles moderately 
prominent as in Fig. 5 

. . Ahstrusomyzus leucocrini (Gillette and Palmer) 

- Cauda usually with fewer than 9 setae; lateral 
abdominal tubercles small or absent (Fig. 18); 
antennal tubercles sometimes strongly pro- 
duced similar to the apterous vivipara (Fig. 6) 

Abstrusomyzus phloxae (Sampson) 



Abstrusomyzus leucocrini (Gillette and 
Palmer 1929), new combination 

(Figs. 4, 5, 9, 13, 17) 

Myzus leucocrini Gillette and Palmer 1929: 
470; Gillette and Palmer 1934: 202; 
Palmer 1952: 338. 

Ovatus leucocrini: Eastop and Hille Ris 
Lambers 1976: 328; Smith and Parron 
1978: 225; Remaudiere and Remaudiere 
1997: 135. 

This species was described from several 
collections made by L.C. Bragg from Fort 
Collins, Colorado, in May of 1916. The 
aphids were found on Leucocrinum mon- 
tanum, a small herb of the Liliaceae. We 
know of no other collections of this aphid 



from Leucocrinum. There are a few indi- 
vidual alate viviparae found in traps and on 
various plants that seem to be this species, 
but lack of good host plant records makes 
any deductions about the biology of this 
species difficult. The presence of many ap- 
terous and alate viviparae on an herb {Leu- 
cocrinum) in May suggests a monoecious 
life cycle. Further collecting will be needed 
to determine whether this species is mo- 
nophagous on Leucocrinum, or feeds on 
other plants as well. 

Within Abstrusomyzus, this species is 
most similar to A. reticulatus and A. valu- 
liae. In the material at hand (19 apterous, 
and 15 alate viviparae), the apterous vivi- 
parae of A. leucocrini almost always have 
partially developed lateral ocelli, and often 
spinal tubercles on the head as well (Fig. 
5). Abstrusomyzus reticulatus lacks both of 
these, and A. valuliae always lacks the lat- 
ter, but usually lacks both. The cauda of A. 
leucocrini is usually more setose, ranging 
from 9-15 setae in both alate and apterous 
viviparae, compared to 4-9 setae on the 
cauda (Figs. 13, 15, 16) of the other two 
species. This species and A. phloxae have 
characteristic brown to orange regions of 
pigmentation surrounding the bases of the 
siphunculi. It is not known whether the oth- 
er two species in the group have this un- 
usual pigmentation pattern. Differences be- 
tween A. leucocrini and A. phloxae are dis- 
cussed under the latter species. Adequate 
descriptions of this species have been pub- 
lished previously (Mason 1940, Palmer 
1952). 

Types of this species are located in the 
USNM. A single paratype slide was also 
obtained on loan from the BMNH, and an- 
other from UCB. Other material examined 
were specimens from the same series as the 
types, some of which were cleared and re- 
mounted for this study. 

Abstrusomyzus phloxae (Sampson 1939), 
new combination 

(Figs. 6, 10, 14, 18) 
Phorodon phloxae Sampson 1939: 174. 
Myzus plantagineus Passerini (misidentifi- 



VOLUME 101. NUMBER 



53 



cation): Williams 1911: 65; Davis 1910: 
495; Davis 1911: 23; Mason 1940: 17. 
Ovatus phloxae: Hille Ris Lambers 1966: 
600; Heie 1972: 450; Eastop and Hille 
Ris Lambers 1976: 329; Smith and Par- 
ron 1978: 225; Remaudiere and Remau- 
diere 1997: 135. 

Abstrusomyzus phloxae was described 
from California based on specimens col- 
lected on Phlox subulata. Sampson de- 
scribed the species in Phorodon Passerini 
under a broad concept of the genus that in- 
cluded species that are today considered 
Ovatus and Myzus. Since Sampson (1939), 
the species has been collected on many oth- 
er plants (Table 1 ). In the eastern half of the 
U.S.A., A. phloxae is most often found on 
the crown, young leaves, and roots of Plan- 
tago major. This led to the misidentification 
of this species by Williams (1911) and Ma- 
son (1940) as Myzus plantagineus Passeri- 
ni. Hille Ris Lambers (1966) was the first 
to understand the identity and wide host 
range of this species. 

This species has been found on many un- 
related host plants, but mostly only in iso- 
lated collections. It has been best studied in 
eastern U.S.A. on Plantago major. It was 
collected on this plant in Illinois, Maryland, 
Nebraska, Pennsylvania, and Virginia in 
May, June, and July of various years. When 
feeding on this plant A. phloxae is often 
tended by ants (Lasius alienus Foerster in 
Maryland) which build earthen "tents" sur- 
rounding the young leaves on which the 
aphids feed. Such "tents" we found in Mar- 
yland housed a mixture of aphids, including 
A. phloxae, Aphis gossypii, and Nearctaphis 
bakeri (Cowen). We reared A. phloxae on 
Plantago on potted plants indoors. To test 
acceptance of two host plants used by other 
Abstrusomyzus species, we allowed the 
population of A. phloxae on Plantago to in- 
crease, and provided Fragaria sp. (the host 
genus of A. valuliae) and Oxalis sp. (the 
host genus of A. reticulatus) in adjacent 
potted plants. Many A. phloxae individuals 
attempted to colonize Fragaria and Oxalis, 



but no colonies were established during the 
two weeks of attempts. The A. phloxae pop- 
ulation eventually killed its host plants. 
This bolstered the somewhat weak morpho- 
logical separation between A. phloxae and 
A. reticulatus and A. valuliae, since despite 
the polyphagy of A. phloxae, it was unable 
to colonize the hosts of these two species. 
The aphids reared on potted plants were 
heavily parasitized by the aphelinid Aphel- 
inus asychis Walker. 

Little is known about the life history of 
this species. It has been collected in every 
month except October and November. In 
Maryland the earliest collection was Feb- 
ruary. In Oregon it is most commonly 
found in August and September. The single 
known collection of oviparae was found in 
Abbotsford, British Columbia on 2 January 
1992. Given that males have never been 
found, and considering that it has been col- 
lected almost throughout the year, we sus- 
pect that A. phloxae is primarily anholo- 
cyclic. Concerted collecting efforts during 
autumn and winter will be required to de- 
termine the life history of this species. 

When A. phloxae is found in nature, it 
almost always lives on the lower leaves of 
its host plant, and generally on plants that 
are low to the ground or have a basal ro- 
sette. When reared on potted plants indoors, 
the aphid thrived on all parts of the plant, 
but moved to the upper parts only when 
populations were very large. This propen- 
sity for living near the ground, but on many 
different plant species, suggests that this 
aphid may be a rare example of a habitat- 
specific, rather than host-specific aphid. 
Other examples of this phenomenon are 
known in the aphids, such as the polypha- 
gous tree feeding aphid Longistigma caryae 
(Harris), which feeds specifically on bark, 
and Rhopalosiphum nymphaeae, which 
feeds on aquatic herbs. 

Chromosome number is often useful as a 
taxonomic character in aphids. For most 
species of Myzus in which it is known, the 
chromosome number is 2n = 12, with a few 
having 2n = 13 or 14. One slide obtained 



54 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



on loan from the BMNH contains a sample 
of apterous viviparae of A. phlo.xae that was 
karyotyped as 2n = 18 by R.L. Blackman. 
Karyotyping of the other Abstrusomyzus 
species may show that chromosome num- 
bers will also support the separation of Abs- 
trusomyzus from similar Myzus-Mke. genera. 

Within Abstrusomyzus, apterous vivipa- 
rae of A. phloxae are easily recognized by 
their pale tergum and large, distinctly con- 
verging antennal tubercles (Fig. 6). Alate 
viviparae are far more difficult to separate 
from other members of the group. The alate 
form of A. reticulatus is unknown. The long 
middle pair of setae on abdominal tergite 
VIII is diagnostic for A. valuliae. This 
leaves A. leucocrini as the primary species 
that may be confused with A. phlo.xae in the 
alate stage. Abstrusomyzus phloxae has rel- 
atively more prominent antennal tubercles, 
and usually has fewer caudal setae; A. 
phloxae normally has eight or fewer caudal 
setae (but sometimes up to 10), compared 
to 9-13 in A. leucocrini (Figs. 13, 14). 
Many A. leucocrini have spinal tubercles on 
the head, which are always absent in A. 
phloxae. Finally, A. phloxae tends to have 
smaller lateral tubercles, and often fainter 
or less extensive abdominal pigmentation 
(Figs. 17, 18). Good descriptions of A. 
phlo.xae can be found in Sampson (1939) 
and Mason (1940). 

Many paratypes of this species were 
seen, having been borrowed from the UCB. 
Other material examined included material 
from all the host plants and states listed for 
this species in Table 1. Seventy eight ap- 
terous viviparae, and 59 alate viviparae 
were seen during this study. 

Abstrusomyzus reticulatus (Heie 1972), 
new combination 

(Figs. 7, 11, 15) 

Ovatus reticulatus Heie 1972: 447; Eastop 
and Hille Ris Lambers 1976: 329; Smith 
and Parron 1978: 225; Remaudiere and 
Remaudiere 1997: 135. 

This species was described based on nine 
specimens collected on O.xalis ?stricta in 



North Carolina. These specimens are still 
the only ones known for the species. They 
include seven apterous viviparae, one ovi- 
para, and one brachypterous male. 

Abstrusomyzus reticulatus is apparently 
monoecious holocyclic based on the occur- 
rence of viviparae, a male, and ovipara to- 
gether on the herbaceous Oxalis. According 
to Heie (1972) the aphids cause the leaves 
of O.xalis to curl. Further collecting will be 
required to determine the host range of this 
species. 

This species can be distinguished from A. 
phlo.xae as described above. Abstrusomyzus 
valuliae is distinct from this species and 
others in the group because of the long se- 
tae on abdominal tergite VIII. Abstruso- 
myzus reticulatus is most similar to A. leu- 
cocrini. These species can be separated 
based on the small number of caudal setae 
in A. reticulatus, its complete lack of spinal 
tubercles on the head, and relatively dark 
antennal segment II. This is the only spe- 
cies of Abstrusomyzus for which the male 
is known. The brachypterous condition of 
the single known male is probably aberrant. 
Heie (1972) provides good descriptions of 
all known morphs. 

Types of this species are housed in the 
USNM (holotype, one paratype), BMNH (3 
apterous vivipara paratypes), C.F Smith 
collection (two apterous viviparae, not 
seen), and in the collection of O.E. Heie 
(one apterous vivipara and one ovipara, not 
seen). 



Abstrusomyzus valuliae (Robinson, 1974), 
new combination 

(Figs. 8, 12, 16) 

Myzus valuliae Robinson 1974: 469. 

Ovatus valuliae: Eastop and Hille Ris 
Lambers 1976: 329; Smith and Parron 
1978: 226; Remaudiere and Remaudiere 
1997: 135. 

Abstrusomyzus valuliae is known only 
from the material used by Robinson for the 
description of the species from Manitoba, 



VOLUME 101, NUMBER 1 



55 



Canada. It was collected several times dur- 
ing the summer of 1973 and once in the 
spring of 1974 on Fragaha vesca, wild 
strawberry. These specimens include many 
apterous and alate viviparae. 

Little is known about the biology of this 
species. It is most likely monoecious, but 
host plant range is unknown. When on Fra- 
gaha, the aphids cause the leaves to curl 
tightly, and the aphids feed inside the curled 
leaves (Robinson 1974). 

This species can be distinguished from A. 
phloxae as described under that species. 
Among the three darkly pigmented species 
of Abstrusomyzus, this species can be most 
easily distinguished by the long middle pair 
of setae on the abdominal tergite VIII. It is 
most similar to A. leucocrini, both species 
frequently having lateral ocelli or traces 
thereof in the apterous vivipara. Robinson 
(1974) described this species thoroughly. 

The holotype of A. valuliae, which we 
did not see, was deposited in the Canadian 
National Collection, Ottawa. Paratypes 
were deposited in the Canadian National 
Collection, BMNH, and the USNM. Many 
paratypes and some additional material 
from the same collections were examined 
for this study. 

Status of Ovatiis 

The removal of four species from Ovatus 
leaves only a single species of the genus in 
North America, O. crataegariiis. This spe- 
cies can be separated from Myzus found in 
North America by its lack of spinules on 
the hind tibia of nymphs, the lack of a dor- 
sal pigmented abdominal patch in the alate 
viviparae, and the presence of many sec- 
ondary rhinaria on antennal segments III, 
IV, and V in alate viviparae. Ovatus cra- 
taegarius can be separated from another 
somewhat similar species, Phorodon hu- 
muli (Schrank), by its first tarsal chaetotaxy 
of 3,3,3, lack of pigmented abdominal patch 
in the alate viviparae, and its antennal tu- 
bercles, which are strongly convergent, but 
lack the extremely prominent finger-like 
process of Phorodon. The primary way that 



Ovatus is separated from Hyalomyzus is the 
swollen siphunculi of the latter genus. An 
excellent diagnosis for the genus Ovatus, as 
it is understood here, is provided by Heie 
(1994) (i.e., Heie's diagnosis does not ad- 
dress the characters of the four species here 
placed in Abstrusomyzus). 

Acknowledgments 

The authors thank Paul A. Brown (The 
Natural History Museum, London), David 
J. Voegtlin (Illinois Natural History Survey, 
Champaign), Cheryl Barr (University of 
California, Berkeley), James LaBonte 
(Oregon State University, Corvallis), and C. 
K. Chan (Agriculture Canada, Vancouver) 
for the loan of specimens. The manuscript 
was improved by the comments of Natalia 
J. Vandenberg and Michael E. Schauff of 
the U.S. Department of Agriculture, Sys- 
tematic Entomology Laboratory, Victor F. 
Eastop of The Natural History Museum, 
London, David J. Voegtlin of the Illinois 
Natural History Survey, and Robert G. 
Foottit of Agriculture Canada, Ottawa. 
Identifications of the ants and aphelinids 
were provided by D. R. Smith and M. E. 
Schauff, respectively, of the U.S. Depart- 
ment of Agriculture, Systematic Entomol- 
ogy Laboratory. 

Literature Cited 

Blackman, R. L. and V. F. Eastop. 1994. Aphids on 
the World's Trees. CAB International, Walling- 
ford, U.K. 987 pp. 

Bremer, K. 1994. Branch support and tree stability. 
Cladistics 10: 295-304. 

Davis, J. J. 1910. A list of the Aphididae of Illinois, 
with notes on some of the species. Journal of Eco- 
nomic Entomology 3: 482-499. 

. 1911. Williams' "The Aphididae of Nebras- 
ka"; a critical review. University Studies (Nebras- 
ka) 11: 253-291 +8 pis. 

Eastop, V. E 1966. A taxonomic study of Australian 
Aphidoidea (Homoptera). Australian Journal of 
Zoology 14: 399-592. 

. 1973. Deductions from the present day host 

plants of aphids and related insects. Symposium 
of the Royal Entomological Society. London 6: 
157-178. 

Eastop, V. F. and D. Hille Ris Lambers. 1976. Survey 



56 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



of the World's Aphids. Dr. W. Junk, b.v.. The 
Hague. 573 pp. 

Gillette, C. P. and M. A. Palmer. 1929. Five New Aphi- 
didae from Colorado. Annals of the Entomologi- 
cal Society of America 22: 468-476 +2 pis. 

. 1934. The Aphidae of Colorado. Part III. An- 
nals of the Entomological Society of America 27: 
133-255. 

Gould, S. J. 1977. Ontogeny and Phylogeny. Harvard 
University Press, Cambridge. 501 pp. 

Heie, O. E. 1972. Ovatus reticulatus, a new species of 
aphid from Oxalis in North Carolina (Homoptera: 
Aphididae). Proceedings of the Entomological So- 
ciety of Washington 74: 446-450. 

. 1994. The Aphidoidea (Hemiptera) of Fen- 

noscandia and Denmark. V. Family Aphididae: 
Part 2 of tribe Macrosiphini of subfamily Aphi- 
dinae. Fauna Entomologica Scandinavica 28. 242 
pp. 

Hille Ris Lambers. D. 1966. Notes on California 
aphids, with descriptions of new genera and new 
species. Hilgardia 37: 569-623. 

Jensen. A. S. 1997. Redefinition of the aphid genus 
Sitobion Mordvilko (Hemiptera: Aphididae) based 
on cladistic analyses, with emphasis on North 
American species. Systematic Entomology 22: 
333-344. 

Maddison, W. P and D. R. Maddison. 1992. Mac- 
Clade: Analysis of Phylogeny and Character Evo- 
lution. Version 3.0. Sinauer Associates, Sunder- 
land. Massachusetts. 

Mason, P. W. 1940. A revision of the North American 



aphids of the genus Myzus. United States Depart- 
ment of Agriculture Miscellaneous Publication 
No. 371: 1-30. 

Miyazaki. M. 1987. Morphology of aphids. pp. 1-25. 
In Minks. A. K. and Harrewijn. P., eds.. Aphids, 
Their Biology. Natural Enemies and Control 2A. 
Elsevier. New York. 

Nielsson, R. J. and D. H. Habeck. 1971. The genus 
Hyalomyzus (Homoptera: Aphididae). with the de- 
scription of a new species. Annals of the Ento- 
mological Society of America 64: 883-887. 

Palmer, M.A. 1952. Aphids of the Rocky Mountain 
Region, Vol. 5. The Thomas Say Foundation. 452 
pp. 

Remaudiere, G. and M. Remaudiere. 1997. Catalogue 
of the World's Aphids. INRA, Paris. 473 pp. 

Robinson, A. G. 1974. Four new species of aphids 
(Homoptera: Aphididae) from Manitoba. Canadi- 
an Entomologist 106: 467-472. 

Sampson, W. W. 1939. California of aphids of the ge- 
nus Phorodon. The Pan-Pacific Entomologist 15: 
173-175. 

Sinith. C. F and C. S. Parron. 1978. An Annotated List 
of Aphididae (Homoptera) of North America. 
North Carolina Agricultural Experiment Station. 
428 pp. 

Swofford. D. L. 1993. PAUP: Phylogenetic Analysis 
Using Parsimony. Version 3.1.1. Computer pro- 
gram distributed by the Illinois Natural History 
Survey. Champaign. Illinois. 

Williams. T. A. 1911. The Aphididae of Nebraska. 
University Studies (Nebraska) 10: 85-175. 



PROC. ENTOMOL. SOC. WASH. 

lOltl), 1999, pp. 57-63 

THE LARVAL INSTARS OF THE WHEAT MIDGE, SITODIPLOSIS 
MOSELLANA (GEHIN) (DIPTERA: CECIDOMYHDAE) 

Raymond J. Gagne and John F. Doane 

(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. De- 
partment of Agriculture, % National Museum of Natural History, MRC 168, Washington, 
DC 20560-0168, U.S.A. (email: rgagne@sel.barc.usda.gov); (JFD) Agriculture and Agri-Food 
Canada Research Centre, 107 Science Place, Saskatoon, Saskatchewan S7H 0X2 Canada. 



Abstract. — The wheat midge, Sitodiplosis mosellana (Gehin), is shown to have three 
larval instars. Each instar is described and illustrated, the initial instar for the first time. 
An alleged synonym of the wheat midge, Cecidomyia amyotii Fitch, is removed from 
synonymy with the wheat midge and considered a dubious name. 

Key Words: wheat midge, Sitodiplosis, larva 



The wheat midge, Sitodiplosis mosellana 
(Gehin), is one of the two most important 
cecidomyiid pests of wheat in North Amer- 
ica, the other being the Hessian fly, Maye- 
tiola destructor (Say). Originally from the 
Palearctic Region, both are now well estab- 
lished in North America. All three larval 
instars of the Hessian fly have been de- 
scribed in detail (Gagne and Hatchett 
1989), but those of the wheat midge are less 
well known, and its first instar has not pre- 
viously been described. 

The general life history of the wheat 
midge or, in the United Kingdom, the or- 
ange wheat blossom midge, has been sum- 
marized by Reeher (1945) and Barnes 
(1956), and more information on certain 
aspects of its attack and feeding were 
treated by Mukerji et al. (1988) and Elliott 
and Mann (1996). Females lay eggs on 
emerging spikes of wheat before anthesis. 
Upon hatching, the larvae crawl to and 
settle upon the developing flower parts, 
where they feed and interfere with the 
proper development of the kernels. The 
full grown larva eventually drops from 
the wheat head, crawls into the soil, and 



constructs a silk cocoon in which the lar- 
va overwinters. 

We show that the wheat midge has three 
larval instars, as do the Hessian fly and 
all other cecidomyiids that have been 
carefully studied (Gagne 1989). Upon 
hatching, the first instar of the wheat 
midge crawls to a feeding site on the de- 
veloping wheat grain, settles, and begins 
to feed. Within two to three days the larva 
molts to the second instar, which contin- 
ues to feed and grow until the third instar 
begins to develop. The second instar skin 
then becomes brittle and serves as a tem- 
porary cocoon for the third instar, which 
does not feed. The fully developed third 
instar can exit from the second instar skin 
immediately but may stay within this skin 
on the kernel for several weeks. Upon 
leaving the temporary cocoon, the third 
instar drops to the ground and burrows 
into the soil where it spins its cocoon. 

In most other cecidomyiids the second 
instar skin is shed as soon as the third in- 
star is fully developed and, in gall-making 
species, can be found crumpled in a com- 
pact mass at the caudal end of the third 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



instar. The situation in the wheat midge is 
somewhat analogous to that of the Hes- 
sian fly in which the second instar skin 
also serves as a cocoon for the non-feed- 
ing third instar, with the difference that 
the third instar of the Hessian fly pupates 
there also. Because the Hessian fly feeds 
head groundwards between the culm and 
leaf sheath, the third instar has to reverse 
position from head to tail within the sec- 
ond instar skin. In this way the pupa of 
the Hessian fly is positioned so that adults 
are able to exit from the wheat sheaths. 
The third instar of the wheat midge is not 
so constrained, so does not need to re- 
verse its position within the second instar 
skin. These two cecidomyiids on grasses 
are not closely related, each belonging to 
a different supertribe, so this rare devel- 
opment of a brittle second instar skin as a 
puparium or larvarium has evidently 
evolved separately. A puparium is present 
also in another grass-infesting gall midge, 
the sorghum midge, Stenodiplosis sorgh- 
icola (Coquillett) (Solinas 1986), which 
belongs to the same supertribe but to a 
different tribe than the wheat midge. A 
puparium is also known for other gall 
midges that do not occur on grasses but is 
still rare. Examples are Thurauia aquatica 
Riibsaamen from sedges in Europe and an 
undescribed species that belongs to no 
known genus from maple seeds in Japan. 
The description and figures that follow 
will allow recognition of each of the three 
instars of the wheat midge. They are nec- 
essary to correct misinformation in Borkent 
(1989) that was based on mixed series of 
three species and in which it was asserted 
that the wheat midge had four instars. 

Methods 

In 1997, collections of wheat heads (cul- 
tivar Roblin) were made by one of us (JFD) 
from a plot area (latitude 52.1951, longi- 
tude 106.1071) near St. Denis, a hamlet 
about 26 miles east of Saskatoon, Saskatch- 
ewan, Canada. Twenty-five primary heads 
were collected at intervals of 1 or 2 days 



from July 7 until July 18 and then at inter- 
vals of 3 to 4 days until July 31. The first 
eggs were found on the outside of the 
glumes on July 7 and the first newly 
hatched larvae on July 9. The larvae were 
removed from the heads and preserved in 
70% alcohol for subsequent examination. 
Some specimens were mounted on micro- 
scope slides using the method outlined in 
Gagne (1989); earlier instars were mounted 
in Hoyer's medium because of their small 
size and the risk of their being lost in the 
various steps involved in balsam mounting. 
Terminology for larval morphology follows 
that in Gagne (1989). 

Description of Larval Instars of 
sitodiplosis mosellana 

The first instar differs markedly from the 
remaining instars. It has only one pair of 
functional spiracles (Figs. 10, 12), which 
are situated on the eighth abdominal seg- 
ment and are relatively large in relation to 
body size compared to the eighth abdomi- 
nal spiracle of other instars (Fig. 13). Its 
cuticle is entirely smooth except for several 
horizontal rows of tiny spicules dorsally 
and ventrally near the anterior part of most 
segments (Figs. 11, 12). The second and 
third instars each have spiracles on the first 
thoracic and on the first through eighth ab- 
dominal segments. The cuticle of the sec- 
ond and third instars is rough, covered al- 
most entirely by raised scale-like bumps 
(Figs. 4-7). The third instar differs from the 
second in having a spatula, the clove- 
shaped dermal structure on the venter of the 
prothorax (Fig. 9). Because this structure 
begins to develop while the third instar is 
still encased in the second instar, the spatula 
is usually visible through the skin of older 
second instars. An additional difference be- 
tween the second and third instars is that 
the two caudalmost terminal papillae are 
more equal in size in the third instar than 
in the second (Fig. 14). A more detailed 
description follows: 

First instar (Figs. 10-12). — Body length 
0.45-1.05 mm. Antenna three times as long 



VOLUME 101, NUMBER 



59 




Figs. 1-7. Sitodiplosis mosellami. 1-3, Brittle second instar skins from which third instar is emerging (1.2) 
■ has emerged (3). 4, Second instar (dorsolateral). 5, Same (lateral). 6, Third instar (dorsal). 7, Same (lateral). 



60 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



as wide. Two pairs of spiracles apparent, 
one on prothoracic segment, the other on 
eighth abdominal segment: prothoracic pair 
evident only as short projection, evidently 
not functional; posterior pair prominent, 
with three acute apical projections. Cuticle 
smooth except for several horizontal rows 
of tiny spicules dorsally and ventrally near 
anterior part of each segment except for 
prothorax. Pattern and number of papillae 
basic for supertribe Cecidomyiidi, their se- 
tae mostly very short, dorsal and pleural pa- 
pillae of eighth segment slightly longer than 
preceding segments, and terminal papillae 
modified as follows: one dorsal, subcaudal 
pair with short setae not surpassing length 
of those on preceding segments; one lateral 
pair with elongate setae several times lon- 
ger than subcaudal pair; two caudal pairs 
with stout, wide setae, the outer of these 
two pairs more than twice length of inner 
pair. 

Second instar (Figs. 4, 5, 8, 9, 14). — 
Body length 1.05-3.30 mm. Antenna about 
twice as long as wide. Spiracles present on 
prothoracic and first through eighth abdom- 
inal segments, eighth abdominal pair slight- 
ly larger than preceding spiracles, none 
with apical projections. Cuticle rugose, cov- 
ered almost completely with raised scale- 
like bumps, these much smaller and ar- 
ranged in several regular anteroventral hor- 
izontal rows on mesothoracic to eighth ab- 
dominal segments. Pattern of papillae sim- 
ilar to that of first instar but setae more con- 
spicuous. Terminal papillae modified as 
follows: two lateralmost papillae with setae, 
the more ventral pair with longer setae; two 
pairs with stout setae, the inner pair with 
setae noticeably thinner than outer pair. 

Third instar (Figs. 6-9, 13, 14). — Body 
length 2.80-3.20 mm. Antenna and spira- 
cles as for second instar. Cuticle also as for 
second instar except for presence of ventral 
prothoracic spatula and horizontal antero- 
ventral rows of bumps more numerous and 
more extensive. Pattern of papillae similar 
to that of second instar. Terminal papillae 
modified as for second instar except inner 




-«9 

Figs. 8, 9. Sitodiplosis mosellana. 8. Third instar 
inside second instar skin (ventral). 9, Detail of same 
(a = spatula). 



pair of caudal, stout setae more nearly equal 
in size to outer pair. 

Remarks. — Borkent (1989) described 
and illustrated four alleged instars for the 
wheat midge, each with a spatula. The first 
instar of that paper is a third instar of an 
undetermined Clinodiplosis sp. Its posterior 
abdominal segment, as illustrated in his Fig. 
2A, has three of the four pairs of terminal 
papillae corniform (short and stout) and 
only one pair setiform, characteristic of spe- 
cies of Clinodiplosis. The presence of a 
spatula indicates that the specimen is a third 
instar. Specimens of these larvae occasion- 
ally are found in association with wheat 
midge larvae. Representatives of this spe- 
cies in the USNM are curated with wheat 
midge larvae until adults are reared and the 
species can be identified further. 

The second instar of Borkent (1989) is a 
third instar of an undescribed species of 
Contahnia (broad sense). It has one pair of 



VOLUME 101. NUMBER 1 



61 




.05 




.03 





dorsal papilla 
pleural papilla 
spiracle 
terminal papillae 



Figs. 10-14. Sitodiplosis mosellana. 10, First instar (dorsal). 1 1, Same, detail of head and first two thoracic 
segments. 12, Same, detail of posterior segments. 13, Third instar, posterior part of eighth segment and terminal 
segment (dorsal). 14, Juxtaposed third instar (upper) and second instar (lower) terminal segments (dorsal). Bar 
lengths are in mm. 



62 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



corniform (short and stout) papillae and 
three pairs of setiform papillae, two of 
which are of similar size. The presence of 
a spatula indicates that this specimen is also 
a third instar Specimens in the USNM of 
similar larvae found in association with the 
wheat midge also have a spatula and are 
definitely third instars. These are also tem- 
porarily curated with wheat midge larvae 
until adults are found that can be identified 
further 

The third and fourth instars of Borkent 
(1989) are actually the second and third in- 
star, respectively, of the wheat midge. Both 
are described in that paper as having a spat- 
ula, but the second instar has none. As not- 
ed above, older specimens of the second in- 
star may appear to have a spatula due to the 
developing third instar inside (Figs. 8, 9). 
One can be certain of the train of instars of 
a particular species by observing nearly ful- 
ly developed preecdysal instars within the 
body of a previous instar, as we have done 
here. 

Borkent (1989) cited Borkent (in press), 
"Description of the larval instars of the 
Wheat Midge Sitodiplosis mosellana (Ge- 
hin) (Diptera: Cecidomyiidae). Can. J. 
Zool." No paper by Borkent on the subject 
of the wheat midge has appeared in the Ca- 
nadian Journal of Zoology. 

This occasion is taken to remove the 
name Cecidomyia amyotii Fitch from syn- 
onymy with the wheat midge. Felt (1925) 
listed C. amyotii as "probably a synonym 
of" the wheat midge and Foote (1965) cat- 
aloged it as a synonym of the wheat midge 
for the first time. According to Fitch's 
(1861) original description of C. amyotii, 
based on three specimens caught at a light, 
the female antenna has "eighteen (?) joints, 
not separated by pedicels, the joints glob- 
ular" The female of the wheat midge has 
12 antennal flagellomeres, each separated 
by a conspicuous pedicel or neck, so C. 
amyotii cannot be the wheat midge. Be- 
cause the types of C. amyotii are presum- 
ably lost and the species cannot be deter- 



mined with certainty, we consider this spe- 
cies a dubious name. 

Acknowledgments 

We thank Nit Malikul for making the 
slide preparations; Cathy Anderson for 
computer assistance in making the plates; 
and, for their comments on drafts of the 
manuscript: Philip S. Barker and Robert J. 
Lamb of Agriculture Canada Research Cen- 
tre, Winnipeg, Manitoba; John W. Brown 
and Allen L. Norrbom of the Systematic 
Entomology Laboratory, USDA, Washing- 
ton, DC; Keith M. Harris, International In- 
stitute of Entomology, CAB, London, UK; 
and Odette Rohfritsch, Institut de Biologic 
Moleculaire des Plantes, Strasbourg, 
France. 

Literature Cited 

Barnes, H. F. 1956. Gall Midges of Economic Impor- 
tance, Vol. 7: Gall Midges of Cereal Crops. Cros- 
by Lockwood & Son, London. 261 pp. and 16 pis. 

Borkent, A. 1989. A review of the wheat blossom 
midge, Sitodiplosis mosellana (Gehin) (Diptera: 
Cecidomyiidae) in Canada. Agriculture Canada 
Research Branch Technical Bulletin 1989-3E. Ot- 
tawa, Ontario. 18 pp. 6 pis. 

Elliott. R. H. and L. W. Mann. 1996. Susceptibility of 
red spring wheat. Triticuin aestivitm L. CV. Ka- 
tepwa. during heading and anthesis to damage by 
wheat midge, Sitodiplosis mosellana (Gehin) 
(Diptera: Cecidomyiidae). The Canadian Ento- 
mologist 128: 367-375. 

Felt, E. P. 1925. Key to gall midges (a resume of stud- 
ies I-VII, Itonididae). New York State Museum 
Bulletin 257: 1-239, pis. 1-8. 

Fitch. A. 1861. Sixth report on the noxious and other 
insects of the State of New York. New York State 
Agricultural Society Transactions (1860) 20: 745- 
868, pis. 1-2. 

Foote. R. H. 1965. Family Cecidomyiidae. //; Stone, 
A., C. W. Sabrosky, W. W. Wirth. R. H. Foote, 
and J. R. Coulson, eds., A Catalog of the Diptera 
of America North of Mexico. U.S. Department of 
Agriculture Handbook 276, 1696 pp. 

Gagne, R. J. 1989. The Plant-Feeding Gall Midges of 
North America. Cornell University Press, Ithaca, 
New York, xiii, 355 pp., 4 pis. 

Gagne, R. J. and J. H. Hatchett. 1989. Larval instars 
of the Hessian fly (Diptera: Cecidomyiidae). An- 
nals of the Entomological Society of America 82: 
73-79. 

Mukerji, M. K.. O. O. Olfert, and J. F Doane. 1988. 



VOLUME 101, NUMBER 1 63 

Development of sampling designs for egg and lar- Northwest. United States Department of Agricul- 

val populations of the wheat midge, Sitodiplosis ture Circular 732: 1-8. 

mosellana (Gehin) (Diptera: Cecidomyiidae), in Solinas, M. 1986. Allocontarinia, n.g., A. sorghicola 

wheat. The Canadian Entomologist 120:497-505. (Coq.) n. comb. (Diptera, Cecidomyiidae). Ento- 

Reeher, M. M. 1945. The wheat midge in the Pacific mologica 21: 23-35. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999, pp. 64-68 

THE LARVA OF PELECINUS POLYTURATOR (DRURY) 
(HYMENOPTERA: PELECINIDAE) 

Norman F. Johnson, Luciana Musetti, James B. Johnson, and Kerry Katovich 

(NFJ, LM) Department of Entomology, The Ohio State University, 1315 Kinnear Road, 
Columbus, OH 43212-1192, U.S.A. (e-mail: Johnson. 2@osu.edu;luciana@iris.biosci. 
ohio-state.edu); (JBJ) Division of Entomology, Department of Plant, Soil, and Entomo- 
logical Sciences, University of Idaho, Moscow, ID 83844-2339, U.S.A. (e-mail: 
djohnson@novell.uidaho.edu); (KK) Department of Entomology, 237 Russell Laborato- 
ries, University of Wisconsin, Madison, WI 53706, U.S.A. 



Abstract. — The first instar larva and exuviae of the last instar larva of Pelecinus po- 
lyturator (Drury) are described, illustrated, and compared with published descriptions of 
other larvae of Proctotrupoidea. 

Key Words: larval morphology, Pelecinidae, Proctotrupoidea 



The application of characters from larval 
morphology to the study of relationships 
within the Hymenoptera is very uneven. 
There are some good examples of the use- 
fulness of immatures as an additional 
source of data (e.g., Evans 1987), but for 
many superfamilies little or nothing is 
known. Progress in this area is hindered for 
several reasons. Most Apocrita are parasit- 
oids of other insects, and the hosts for a 
number of groups are very poorly known. 
Many species are internal parasitoids; thus 
it is difficult to obtain early instars and of- 
ten the larvae are highly simplified in struc- 
ture. 

Within the Hymenoptera a remarkable ar- 
ray of larval body plans is found. Clausen 
(1940) outlined fourteen types among the 
parasitic Hymenoptera alone. Some of these 
are highly simplified, "embryonic," forms, 
while others possess well-developed, ex- 
aggerated mandibles and caudal appendag- 
es. Later instars usually converge upon a 
generalized, hymenopteriform type. Little 
phylogenetic pattern has been found amidst 
this variety; at present we are confronted 



with a diversity of forms without any un- 
derlying organizing principle. 

The superfamily Proctotrupoidea s. str. is 
comprised of ten extant families of internal 
parasitoids. The hosts for three of these 
(Renyxidae, Austroniidae, and Peradeni- 
idae) are unknown, and only a bare mini- 
mum of information is available for the 
Monomachidae and Roproniidae. The only 
families for which immature stages have 
been described are the Diapriidae (a large 
group of nearly 2,000 recognized species), 
Proctotrupidae (331 described species), and 
Heloridae (a relict group of 10 extant spe- 
cies). In total, the larvae of only six species 
from this complex have been described. We 
report here on the larvae of another family, 
the Pelecinidae, a small group (only one 
species currently recognized) of uncertain 
affinities (Rasnitsyn 1980, Dowton et al. 
1997). 

Materials and Methods 

Five parasitoid larvae were dissected 
from larvae of Scarabaeidae (Coleoptera), 
and preserved in ethanol. The specimens 



VOLUME 101, NUMBER 1 



65 



were found in the posterior two-thirds of 
the abdomen of the host. Three final instar 
exuviae were found attached to scarab re- 
mains from which Pelecinus had pupated. 
Specimens are stored in the collections of 
JBJ; the Ohio State University; the Insect 
Research Collection, University of Wiscon- 
sin; and El Colegio de la Frontera Sur, San 
Cristobal de las Casas, Chiapas. Illustra- 
tions were made using a camera lucida of 
whole specimens under alcohol and exuviae 
in temporary slide mounts in glycerine jel- 

ly- 

Pelecinus polyturator (Drury) 

Material examined. — USA. Michigan, 
Newaygo Co., 18 April 1974, host in soil 
of oak forest, ex Phyllophaga, one first in- 
star; Branch Co., 23 May 1974, hosts in soil 
of oak-hickory forest, four first instars, 
three from large 5 cm long larvae, probably 
Phyllophaga, one from small 2.5 cm larva, 
possibly Serica sp. Wisconsin, Marquette 
Co., 11 August 1992, in sandy soil of forest 
meadow, ex larva of Phyllophaga drakei 
(Kirby) one final instar exuviae; Jackson 
Co., 4 June 1992, ex P. drakei, one final 
instar exuviae; Oconto Co., 28 May 1996, 
ex Phyllophaga rugosa (Melsheimer), one 
final instar exuviae. MEXICO. Chiapas, Te- 
nejapa, Balun Canal, 2,300 m, 14 February 
1997, ex Phyllophaga obsoleta (Blanchard) 
third instar, one final instar exuviae. 

First instar (Figs. 1-4). — Length 3.3-5.3 
mm; mandibulate larva (Clausen 1940); 
head capsule well-developed, covering dor- 
sal and lateral sides of head, margins darkly 
pigmented, sclerotization extending beyond 
margins, gradually disappearing posterior- 
ly; epicranial suture (Fig. 2, es) well-devel- 
oped; no indication of eyes; antenna (Figs. 
1-3, a) indicated by small paired submedial 
papilla; clypeolabral area (Fig. 3, cl) largely 
membranous, supported by ovoid sclero- 
tized ring, dorsal portion of this ring some- 
times incomplete; labrum with two medial 
tubercles (Fig. 1, It); mandible (Figs. 1, 3, 
md) strongly developed, falcate, bearing a 
small subapical tooth; maxilla (Fig. 1, 3, 



mx) supported anteriorly by narrow stipital 
sclerite, otherwise lobelike, membranous; 
maxillary palp, labium, and labial palp un- 
differentiated; head supported internally by 
extensive, strongly pigmented tentorium 
(Fig. 4, tn) in shape of central plate with 
anterior extensions continuous with labral 
sclerite, lateral arms surrounding base of 
mandibles, and broad posterior bilobed 
plate in labial region, a central ovoid fora- 
men visible, anterior to this with more 
strongly pigmented triangular prominence, 
anterior apex of triangle produced into 
small costa extending into labrum; no pro- 
legs visible; body with indeterminate num- 
ber of segments, without setae, apex of ab- 
domen acute; no spiracles visible. 

Final instar (Fig. 5). — Head capsule with 
posterior sclerotized, pigmented band, oth- 
erwise largely membranous; mandible (md) 
very small, weakly articulated with head; 
antenna, labrum, maxilla, maxillary palp in- 
distinguishable; labium (lb) visible as me- 
dial triangular raised surface behind man- 
dibles, with large circular field correspond- 
ing to each labial palp (//?), a small central 
area presumably representing opening of la- 
bial gland (Ig) between palpi; mouthparts 
unsupported by sclerotized pleurostoma or 
hypostoma; body with 7 pairs of spiracles 
visible; tracheae well-developed. 

Discussion 

The exuviae of the last instar larvae are 
associated with pharate adult Pelecinus po- 
lyturator, and their identity is unequivocal. 
The early instar larvae, however, are strik- 
ingly divergent in structure from the exu- 
viae. Our determination of them was based 
on the fact that they were internal parasit- 
oids dissected from larvae of Phyllophaga 
Harris (Coleoptera: Scarabaeidae), the only 
recorded host in the United States and Can- 
ada. The specimens also were collected in 
an area in which Pelecinus was very abun- 
dant. Muesebeck (1979) recorded Tiphia 
berbereti Allen, T. tegulina Malloch, T. 
transversa Say, T. vulgaris Robertson, and 
T. intermedia Malloch (Tiphiidae); Myzin- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs 1-5 Pelecinus poh'turator, larva. 1-4, Head of first instar 1. Lateral view. 2, Dorsal view. 3. Frontal 
view, specimen with mandibles closed. 4, Frontal view, specimen with open mandibles exposmg tentonum. 5 
Mouthparts from final instar exuviae, right mandible detached. Abbreviations: a = antenna; cl - clypeolabral 
area; es = epicranial suture; lb = labium; Ig = opening of labial gland; Ip = labial palp; It = labral tubercle; 
md = mandible; mx = maxilla; tn = tentorium. Scale in mm. 



VOLUME 101, NUMBER 1 



67 



um quinquecinctiim (Fabricius) (Tiphiidae); 
and Ophion nigrovarius Provancher 
(Ichneumonidae) as parasitoids of Phyllo- 
phaga. Woodruff and Beck (1989) listed a 
second species of Ophion as well as a num- 
ber of additional species of tiphiids and sco- 
liids. We ruled out the aculeates because 
they are external parasitoids. Ichneumo- 
noids usually are characterized by the pos- 
session of a hypostomal spur (Short 1978), 
a structure that was not observed in these 
specimens. 

Determination of the number of larval in- 
stars of internal parasitoids requires large 
numbers of observations of cohorts of 
known age in order to detect structural 
changes associated with molting. This has 
not been done yet for any species of proc- 
totrupoid, and no one has yet been able to 
rear Pelecinus through its life cycle. We 
could not determine the age of the observed 
larvae directly or infer their age from pub- 
lished observations of related species. Clau- 
sen (1940) stated that the characteristics 
that set apart mandibulate larvae are lost at 
the first molt. This was confirmed by Clan- 
cy (1946) in his studies of Helorus, another 
proctotrupoid. Therefore, we concluded that 
the larvae dissected from the hosts must be 
late first instars. 

Very little information on the immature 
stages of proctotrupoids exists to form a 
context in which to discuss the structural 
features of Pelecinus. Larvae have been de- 
scribed and illustrated for Helorus anom- 
alipes (Panzer) (Heloridae; Clancy 1946), 
an unidentified species of Basalys West- 
wood (Diapriidae; Simmonds 1952), Basa- 
lys tritoma Thomson (Diapriidae; Wright et 
al. 1946), Coptera silvesthi (Kieffer) (Dia- 
priidae; Pemberton and Willard 1918), Par- 
acodrus apterogynus (Haliday) (Proctotru- 
pidae; Zolk 1924), Phaenoserphus viator 
(Haliday) (Proctotrupidae; Eastham 1929), 
and Brachyserphus parvulus (Nees ab 
Esenbeck) (Proctotrupidae; Osborne 1960). 
Large, sickle-shaped mandibles have been 
reported in the first instar for all these spe- 
cies. A sclerotized head capsule was de- 



scribed in H. anomalipes, B. tritoma, and 
C silvestril. Phaenoserphus viator lacks a 
coinplete head capsule, but does have a 
sclerotized ring surrounding the mouth- 
parts. Distinct antennal lobes are found in 
the helorid, Basalys spp. and the proctotru- 
pids, larger and more prominent than the 
structures found in Pelecinus. We observed 
no prolegs on any of the first instar larvae; 
these structures have been reported for H. 
anomalipes, P. viator, B. parvulus, and an 
unidentified proctotrupid (Clausen 1940; 
presumably Nothoserphus scymni Ash- 
mead). Two of our specimens have paired, 
nipple-like protuberances beneath the pos- 
terior portion of the head capsule (visible 
in Fig. 1). Because of their position, we 
hesitate to call these prolegs or to homol- 
ogize them with the labial palpi. The num- 
ber of observed spiracles reported varies 
from three {B. tritoma, C. silvestrii) to ten 
pairs {P. viator). 

The most striking feature we observed in 
the first-instar larva was the large tentorial 
endoskeleton. A similar structure was very 
briefly described in P. viator by Eastham 
(1929), suggesting that it may not have 
been as apparent or strongly pigmented as 
in Pelecinus. The tentorium is not men- 
tioned in the other descriptions. 

The larval specimens were dissected 
from hosts in the spring (18 April, 1974; 23 
May, 1974) in Michigan. Therefore, it ap- 
pears that the species overwinters as late 
first instars within the Phyllophaga larvae. 
No more than a single larva was found in 
any one host. Three specimens were recov- 
ered from large (5 cm) hosts, presumably 
the final instar of the beetle. A fourth was 
found in a much smaller larva, either a 
much younger specimen or a different ge- 
nus, perhaps Serica MacLeay (Coleoptera: 
Scarabaeidae). Host size may contribute to 
the large variation in size of adult Peleci- 
nus. 

Acknowledgments 

Thanks to Lorena Ruiz-Montoya (San 
Cristobal de las Casas, Mexico) and Daniel 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



K. Young (Madison, WI) for the loan of 
specimens. This material is based in part 
upon work supported by the National Sci- 
ence Foundation under Grant No. DEB- 
9521648. 

Literature Cited 

Clancy, D. W. 1946. The insect parasites of the Chry- 
sopidae (Neuroptera). University of California 
Publications in Entomology 7: 403-496. 

Clausen, C. P. 1940. Entomophagous Insects. Mc- 
Graw-Hill Book Company, Inc., New York. 688 
pp. 

Dowton, M., A. D. Austin, N. Dillon, and E. Bar- 
towsky. 1997. Molecular phylogeny of the apo- 
critan wasps: the Proctotrupomorpha and Eva- 
niomorpha. Systematic Entomology 22: 245—255. 

Eastham, L. E. S. 1929. The post-embryonic devel- 
opment of Phaenoserphiis viator Hal. (Proctotry- 
poidea), a parasite of the larva of Pterostichus ni- 
ger (Carabidae), with notes on the anatomy of the 
larva. Parasitology 21: 1-21. 

Evans, H. E. 1987. Order Hymenoptera, pp. 597-710. 
//; Stehr. F. W., ed.. Immature Insects, Vol. 1. Ken- 
dall/Hunt Publishing Company, Dubuque, Iowa. 
754 pp. 

Muesebeck, C. E W. 1979. Pelecinoidea, pp. 1119- 
1120. //; Ki-ombein, K. V., P D. Hurd, Jr, D. R. 
Smith, and B. D. Burks, eds. Catalog of Hyme- 
noptera in America north of Mexico. Smithsonian 



Institution Press, Washington, DC. Vol. 1, 1198 
pp. 

Osborne, P. 1960. Observations on the natural enemies 
of Meligethes aeneus (F.) and M. viridescens (F.) 
[Coleoptera: Nitidulidae]. Parasitology 50: 91- 
110. 

Pemberton, C. E. and H. F Willard. 1918. A contri- 
bution to the biology of fruit-fly parasites in Ha- 
waii. Journal of Agricultural Research 15: 419- 
466. 

Rasnitsyn, A. P. 1980. [The origin and evolution of the 
Hymenoptera.] Trudy Paleontologicheskogo Insti- 
tuta 174: 1-190. 

Simmonds, F J. 1952. Parasites of the frit-fly, C>.sr/- 
nella frit (L.), in eastern North America. Bulletin 
of Entomological Research 43: 503-542. 

Short, J. R. T 1978. The final larval instars of the 
Ichneumonidae. Memoirs of the American Ento- 
mological Institute No. 25, 508 pp. 

Woodruff, R. E. and B. M. Beck. 1989. The scarab 
beetles of Florida (Coleoptera: Scarabaeidae). Part 
II. The May or June beetles (genus Pliyllophaga). 
Arthropods of Florida and Neighboring Land Ar- 
eas 13. 226 pp. 

Wright, D. W., Q. A. Geering, and D. G. Ashby. 1946. 
The insect parasites of the carrot fly, Psila rosae. 
Fab. Bulletin of Entomological Research 37: 507- 
529. 

Zolk, K. 1924. Pciracodrus apterogymis Halid. biolo- 
gia kohta. Zur Biologic von Poracodrus aptero- 
gynus Halid. Tartu Ulikooli Entomoloogia-Katse- 
jaama Teadaanded 5: 3-10. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999. pp. 69-74 

NEW GENUS AND NEW SPECIES OF AMORBINI (HETEROPTERA: 
COREIDAE) FROM AUSTRALIA 

Harry Brailovsky and Gerasimos Cassis 

(HB) Departamento de Zoologia, Institute de Biologia UNAM, Apdo Postal No. 70- 
153, Mexico, 04510 D.F., Mexico (e-mail: coreidae@servidor.unam.mx); (GC) Australian 
Museum, 6' 8 College Street, PO. Box A285, Sydney South, N.S.W. 2000, Australia. 



Abstract. — Kormijirania, n. gen., and two new species, K. magna and K. parva, col- 
lected in Eastern Australia are described in the tribe Amorbini (Coreidae). Habitus illus- 
trations, drawings of the male and female genitalia, and a key are provided. 

Key Words: Insecta, Heteroptera, Coreidae, Amorbini, new genus, new species. Eastern 
Australia 



The tribe Amorbini (Hemiptera: Heter- 
optera: Coreidae) is represented by 7 genera 
in the Australian region: Acroelytron Mayr 
(1 species), Amorbus Dallas (15 species), 
Crieius Stal ( 1 species), Gelonus Stal ( 1 
species), Kurnaina Distant (1 species), 
Tambourina Distant (1 species), and a new 
genus (2 species) (Brailovsky and Mon- 
teith, in press). 

The present paper adds one new genus 
and two new species of Amorbini from 
eastern Australia. For several years, the spe- 
cies discussed here have remained unnamed 
and undescribed in the hope that additional 
specimens would be found. However, it 
seems desirable to draw the attention of en- 
tomologists to these species in the hope that 
additional specimens will be collected and 
some information obtained on their ecology 
and food requirements. Two striking fea- 
tures of this new genus are the upturned 
juga forming a stout long horn or conical 
tubercle, and the mandibular plate expand- 
ed on a remarkable stout conical tubercle. 

The following abbreviations are used for 
the institutions cited in this paper: BPBM 
(Bernice P. Bishop Museum, Honolulu, Ha- 
waii); SAMA (South Australian Museum, 



Adelaide); UNAM (Instituto de Biologia, 
Universidad Nacional Autonoma de Mexi- 
co). 

All measurements are given in millimeters. 

Kormijirania Brailovsky and Cassis, 
new genus 

Diagnosis. — This new genus resembles 
Gelonus Stal (1865) in having the head 
wider than long, tylus unarmed and extend- 
ing anteriorly to the juga, antenniferous tu- 
bercles unarmed, tibiae sulcate, not foliate, 
and abdominal sternite VII of the female 
with plica and fissura. It is easily distin- 
guished because it is the only known genus 
in the tribe Amorbini with the upturned 
juga forming a stout long horn or conical 
tubercle, and the mandibular plate striking- 
ly expanded on a large conical tubercle. 

Kormijirania has a stout antennal seg- 
ment I, short, and barely crested, antennal 
segments II and III almost cylindrical, bare- 
ly flattened, buccula anteriorly with a clear 
spine-like projection, pronotum slightly 
wider than long, and fore and middle fem- 
ora ventrally with small granules or tiny 
spine-like projections, never with a large 
and laminate subdistal spine. In Gelonus the 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



juga are flattened, the mandibular plate un- 
armed, antennal segments I to III uniformly 
cylindrical, not crested or flattened, buccula 
anteriorly rounded, pronotum clearly wider 
than long, and fore and middle femora ven- 
trally provided with a large and unique sub- 
distal laminate spine. 

A new genus (Brailovsky and Monteith, 
in press), is related to Gelonus and Konni- 
jirania and is recognized by the laterally 
compressed tylus, which is projected up- 
ward as an acute projection, with femora 
unarmed, juga flat, mandibular plate un- 
armed, and antennal segments I to III cylin- 
drical. 

Description. — Macropterous, body 
stout, moderate sized. Head: Width 
across eyes greater than length, quadrate, 
dorsally flat, non declivent, barely pro- 
duced beyond antenniferous tubercles, 
with deep circular pit close to base of ty- 
lus; tylus unarmed, slightly deflexed; juga 
shorter than tylus, upturned to form a 
stout long horn or conical tubercle; anten- 
niferous tubercle unarmed, prominently 
produced, wide, separated by distance 
greater than their own width; sides of 
head in front of eyes almost straight; an- 
tennal segment I robust, thickest, barely 
crested, constricted basally; segments II 
and III stout, cylindrical, barely flattened; 
segment IV fusiform; antennal segment II 
longest, IV shortest, and III longer than I; 
ocelli conspicuous, closer to eyes, and lo- 
cated on an hypothetical line with lower 
margin slightly above lower margin of 
eyes; preocellar pit deep; eyes protruding; 
postocular tubercle markedly produced; 
buccula rounded, short, not projecting be- 
yond anterior third of antenniferous tu- 
bercles, slightly raised, anteriorly with a 
sharp spine-like projection, and posteri- 
orly closed; rostrum reaching posterior 
third of mesosternum or anterior third of 
metasternum; rostral segment III shortest, 
I longest, and II longer than IV; mandib- 
ular plate expanded on a strong conical 
tubercle or acute tooth; ventrally with a 
deep longitudinal groove along midline to 



receive first and anterior third of rostral 
segment II; mandibular plate with mark- 
edly stout conical tubercle. 

Thorax: Pronotum wider than long, tra- 
peziform, non declivent; collar wide; fron- 
tal angles produced forward as rounded 
teeth; humeral angles rounded, not exposed; 
anterolateral margins slightly emarginate, 
obliquely straight, finely serrate; posterolat- 
eral margins sinuate, entire; posterior bor- 
der concave, entire; callar region indistinct, 
transversely flat, separated along midline by 
an obscure longitudinal groove; posterior 
lobe with transverse ridge, distinctly raised; 
prosternum mesally with a slight depres- 
sion; mesosternum with deep longitudinal 
groove to receive rostrum; anterior third of 
mesosternum without lateral expansions; 
anterior lobe of metathoracic peritreme el- 
evated, iiTegularly reniform, posterior lobe 
sharp, small. 

Legs: Short; femora slightly incrassate; 
fore and middle femora densely granulate, 
with small spine-like projections both dor- 
sally and ventrally; hind femur slightly 
granulate, ventrally armed with two short 
subdistal spines or very tiny tubercles; tib- 
iae terete, sulcate. 

Scutellum: Triangular, longer than wide; 
apex truncated or subacute; disc flat. 

Hemelytra: Macropterous, almost reach- 
ing apex of last abdominal segment; costal 
margin emarginate; apical margin straight; 
apical angle short, barely reaching middle 
third of hemelytral membrane. 

Abdomen: Connexival segments reflexed 
above margin of hemelytron at rest; upper 
margin weakly senate; posterior angles of 
connexival segments simple, not spinose; 
abdominal spiracles II to VII submarginal, 
closer to middle third. 

Male genitalia: Genital capsule: Simple; 
posteroventral border slightly concave, with 
posterolateral angles broadly produced 
(Figs. 1, 2). 

Female genitalia: Abdominal sternite 
VII with plica and fissura; plica triangular, 
slightly elevated, apically subacute, and 
reaching anterior third of sternite VII. 



VOLUME 101, NUMBER 1 



71 







Figs. 1-4. 1, 2. Male genital capsule of Kormijirania par\a. 1, Caudal view. 2, Lateral view. 3, 4, Female 
genital plates of K. magna. 3, Caudal view. 4, Lateral view. 



Genital segments: Gonocoxae I enlarged 
dorso-ventrally, in caudal view closed, in 
lateral view slightly convex, with upper 
margin sinuate; paratergite VIII triangu- 
lar, with spiracle visible; paratergite IX 
square, larger than paratergite VIII (Figs. 
3,4). 

Integument: Body surface rather dull, al- 
most glabrous. Head, antennal segments I 
to III, pronotum, clavus, corium, legs, con- 
nexival segments, propleuron, mesopleu- 
ron, metapleuron, and pleural abdominal 
sterna densely granulate; prostemum, me- 
sosternum, metasternum, abdominal sterna, 
and genital plates almost smooth; posterior 
lobe of pronotum, clavus and corium, 
densely punctate. 

Etymology. — We are pleased to name 
this new genus for Dr. Nicholas A. Kor- 
milev, distinguished hemipterist. 

Type species. — Kormijirania parva Brai- 
lovsky and Cassis, new species. 



Kormijirania magna Brailovsky and 
Cassis, new species 

(Figs. 3-5) 

Description. — Measurements: Female: 
Head length 1.39; width across eyes 2.04; 
interocular space 1.30; interocellar space 
0.46; preocular distance 1.05; length of an- 
tennal segments: I, 1.79; II, 2.13; III and IV 
mutilated. Pronotum: Total length 2.54, 
width across frontal angles 1.89; width 
across humeral angles 2.97. Scutellar length 
1.51; width 1.24. Total body length 12.10. 

Female: Dorsal coloration: Head, anten- 
nal segments I and II (III and IV mutilated), 
anterior lobe of pronotal disk, clavus and 
corium dirty yellow with red-brown to 
chestnut-orange punctures and tubercles; 
posterior lobe of pronotal disc dirty yellow 
with black to red-brown punctures; scutel- 
lum dark brown to black; apical angle of 
corium almost black; hemelytral membrane 
dirty white, with veins, basal angle, and 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 5. Dorsal view of Kormijirania magna (9). 



VOLUME 101. NUMBER 



73 




Fig. 6. Dorsal view of Kormijirania pan a (<5). 



scattered spots pale brown; connexival seg- rostral segments I to IV (apex of IV black). 



ments III to VIII light orange yellow, with 
upper margin of posterior third, and tuber- 
cles brown; connexival segment IX pale or- 
ange yellow, with black quadrate spot on 



and legs dirty yellow with red-brown tu- 
bercles; abdominal sterna, and genital 
plates shiny yellow, with pale orange dis- 
coidal spots. Structure: Genital plates as in 



middle third. Ventral coloration: Including Figs. 3, 4. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Male: Unknown 

Type material. — Holotype: 9, Australia, 
Queensland, Mackay (without additional 
data) (BPBM). 

Etymology. — Named for its large size; 
from the Latin word ma gnus. 

Kormijirania parva Brailovsky and 
Cassis, new species 

(Figs. 1, 2, 6) 

Description. — Measurements: Male: 
Head length 1.20; width across eyes 1.86; 
interocular space 1.17; interocellar space 
0.43; preocular distance 0.96; length of an- 
tennal segments: I, 1.24; II, 1.76; III, 1.39; 
IV, 1.14. Pronotum: Total length 1.70; 
width across frontal angles 1.51; width 
across humeral angles 1.96. Scutellar length 
0.98; width 0.80. Total body length 9.00. 
Female: Head length 1.35; width across 
eyes 1.92; interocular space 1.20; interocel- 
lar space 0.43; preocular distance 0.96; 
length of antennal segments: I, 1.36; II, 
1.87; III, 1.54; IV, mutilated. Pronotum: To- 
tal length 2.10; width across frontal angles 
1.70; width across humeral angles 2.25. 
Scutellar length 1.24; width 1.02. Total 
body length 10.00. 

Male: Dorsal coloration: Yellow with 
punctures orange hazel; ocellar tubercle red 
brown; antennal segments I to III yellow 
with red brown tubercles; segment IV with 
anterior half whitish yellow, and posterior 
half orange hazel; humeral angles red 
brown; apex of scutellum whitish yellow; 
hemelytral membrane dirty white, with 
veins, basal angle, and scattered spots pale 
brown; connexival segments II to VII yel- 
low with upper margin of posterior third or 
entirely the posterior third reddish brown; 
abdominal segments I to VI light orange 



yellow and VII yellow with H-shaped dark 
spot. Ventral coloration: Included rostral 
segments I to IV (apex of IV black), buc- 
cula, and anterior and posterior lobe of 
metathoracic peritreme light yellow with 
some red-brown tubercles; genital capsule 
dark brown with three irregular yellow 
spots; legs yellow with red to pink tuber- 
cles. Structure: Genital capsule as in Figs. 
1-2. 

Female: Similar to male. 

Type material. — Holotype: 6, Australia, 
Cairns District, Col. A. M. Lea (without ad- 
ditional data) (SAMA). Paratope: 1 9, 
same data as holotype (SAMA). Both spec- 
imens glued on the same card. 

Etymology. — Named for its small size; 
from the Latin word pana, rather small. 

Key TO Species of Kormijirania 

I . Femora dirty yellow without red to pink tuber- 
cles (Fig. 5): antennal segment I longer than 

1.70, and II longer than 2.00 

magna Brailovsky and Cassis, new species 

- Femora yellow with red to pink tubercles (Fig. 
6); antennal segment I shorter than 1.40, and 

II shorter than 1 .90 

pana Brailovsky and Cassis, new species 

Acknowledgments 

We thank Gordon F Gross (SAMA) and 
Gordon M. Nishida (BPBM) for the loan of 
specimens, and Ernesto Barrera (UN AM), 
Albino Luna (UNAM), and Jesus Contreras 
for preparation of dorsal view illustrations 
and male and female genitalia. 

Literature Cited 

Brailovsky, H. and G. B. Monteith. In press. A new 
genus of Amorbini ( Heteroptera: Coreidae) from 
Australia, with two new species. Memoirs of the 
Queensland Museum. 

Stal. C. 1865. Hemiptera Africana II. Stockholm: 
Nordstedtiana. 200 pp. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 75-85 

LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF 

TRUPANEA ARIZONENSIS MALLOCH (DIPTERA: TEPHRITIDAE) ON 

TRIXIS CALIFORNICA KELLOGG VAR. CALIFORNICA (ASTERACEAE) IN 

SOUTHERN CALIFORNIA 

Richard D. Goeden and Jeffrey A. Teerink 

Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- 
mail: rgoeden@ucracl.ucr.edu) 



Abstract. — Trupanea arizoneiisis Malloch is a monophagous, univoltine fruit fly (Dip- 
tera: Tephritidae) infesting flower heads of Trixis californica Kellogg var. californica 
(Asteraceae) in southern California. The egg, first- and third-instar larvae, and puparia 
are described and figured for the first time. As with each of the eight other Trupanea 
species previously studied, the lateral spiracular complex of the third instar is unique to 
T. arizonensis, comprising three veiTuciform sensilla on the meso- and metathorax and 
two verruciform sensilla on each abdominal segment. The third instar of T. arizonensis 
also differs from those of other Trupanea species previously studied in that both the meso- 
and metathorax are circumscribed by verruciform sensilla. The life cycle is of the aggre- 
gative type; whereby, the adults are long-lived and comprise the over-summering and 
over-wintering stage that returns with winter rainfall to aggregate on regrowing host plants 
for mating, and for oviposition during spring in the preblossom flower heads. The first 
instar tunneled into a single floral tube or ovule of a single immature floret, and each 
second and third instar continued its development by feeding principally on sap conducted 
to the excavated distal end of the same, then slightly stunted ovule. Pupariation occurs in 
the open flower heads, within which an average of 13% of the achenes were destroyed 
among heads containing puparia. Pteromahis sp. (Hymenoptera: Pteromalidae) was reared 
from individual puparia of T. arizonensis as a solitary, larval-pupal endoparasitoid. 

Key Words: Insecta, Trupanea, Asteraceae, nonfrugivorous Tephritidae, biology, tax- 
onomy of immature stages, flower-head feeding, monophagy, host-plant 
range, parasitoid 



Trupanea arizonensis Malloch (Diptera: species from southern California (Cavender 

Tephritidae) is a rarely collected, monoph- and Goeden 1982; Goeden 1987, 1988; 

agous or nearly monophagous species be- Goeden and Teerink 1997, 1998; Goeden et 

longing to one of the larger and more wide- al. 1998a, b; Headrick and Goeden 1991; 

spread genera of nonfrugivorous fruit flies Knio et al. 1996b), along with descriptions 

in North America and California (Foote and of the immature stages of eight of these 

Blanc 1963, Foote et al. 1993). But, being species (Cavender and Goeden 1982, Goe- 

of little or no economic importance, most den and Teerink 1997, 1998; Goeden et al. 

species of Trupanea remained little known 1998a, b; Headrick and Goeden 1991, Knio 

(Foote 1960, Foote et al. 1993) until de- et al. 1996a, Teerink and Goeden 1998). 

tailed life histories were published for nine This paper describes the life history and 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



most immature stages of a tenth species, T. 
arizonensis. 

Materials and Methods 

This study was based in large part on dis- 
sections of subsamples of flower heads of 
Trixis californica Kellogg var. californica 
(Asteraceae) infested by T. arizonensis 
from samples collected during 1991-1995 
in the low-elevation Colorado (northern 
Sonoran) Desert and high-elevation, eastern 
Mojave Desert in southern California in the 
manner described by Goeden (1985, 1992). 
The principal study sites in the Colorado 
Desert were Valliceto Valley at the mouth 
of Smugglers Canyon, 440-m elevation, 
San Diego Co.; the Edmund C. Jaeger Na- 
ture Preserve at 847 m. Desert Center, Riv- 
erside Co.; and Chino Canyon, 1 km NW 
of Palm Springs, at 270 m. Riverside Co. 
One-liter samples of excised, immature and 
mature flower heads containing eggs, lar- 
vae, and puparia were transported in cold- 
chests in an air-conditioned vehicle to the 
laboratory and stored under refrigeration 
for subsequent dissection, photography, de- 
scription, and measurement. Nine eggs, 18 
first- and 16 third-instar larvae, and six pu- 
paria dissected from flower heads were pre- 
served in 70% EtOH for scanning electron 
microscopy (SEM). Additional puparia 
were placed in separate, glass shell vials 
stoppered with absorbant cotton and held in 
humidity chambers at room temperature for 
adult and parasitoid emergence. Specimens 
for SEM were hydrated to distilled water in 
a decreasing series of acidulated EtOH. 
They were osmicated for 24 h, dehydrated 
through an increasing series of acidulated 
EtOH and two, 1-h immersions in Hexa- 
methlydisilazane (HMDS), mounted on 
stubs, sputter-coated with a gold-palladium 
alloy, and studied with a Philips XL30-FEG 
scanning electron microscope in the Insti- 
tute of Geophysics and Planetary Physics, 
University of California, Riverside. 

Most adults reared from isolated puparia 
were individually caged in 850-ml, clear- 
plastic, screened-top cages with a cotton 



wick and basal water reservoir and provi- 
sioned with a strip of paper toweling im- 
pregnated with yeast hydrolyzate and su- 
crose. These cages were used for longevity 
studies in the insectary of the Department 
of Entomology, University of California, 
Riverside, at 25 ± 1°C, and 14/10 (L/D) 
photoperiod. 

Plant names used in this paper follow 
Hickman (1993) and Bremer (1994); te- 
phritid names and adult terminology follow 
Foote et al. (1993). Terminology and tele- 
graphic format used to describe the imma- 
ture stages follow Knio et al. (1996a), Goe- 
den and Teerink (1997, 1998), Goeden et 
al. (1998a, b), and Teerink and Goeden 
(1998), and our earlier works cited therein. 
Means ± SE are used throughout this paper. 
Voucher specimens of T. arizonensis and its 
parasitoids reside in the research collections 
of RDG; preserved specimens of eggs, lar- 
vae and puparia are stored in a separate col- 
lection of immature Tephritidae acquired by 
JAT and now maintained by RDG. 

Results and Discussion 

Taxonomy 

Adult. — Trupanea arizonensis was first 
described by Malloch (1942) as Trypanea 
arizonensis. Malloch (1942), Foote (1960), 
Foote and Blanc (1963), and Foote et al. 
(1993) pictured the wing pattern, which 
Foote (1960) described as not sexually di- 
morphic, but essentially alike in every im- 
portant character in the female and male. 
However, the variable nature of this wing 
pattern, especially among males, initially 
caused Goeden and Ricker (1989) to mis- 
identify specimens reared from Trixis cali- 
fornica as Trupanea actinobola (Loew), an- 
other variable species (Foote and Blanc 
1963, Foote et al. 1993, Goeden et al. 
1998b). Goeden (1992) corrected this error 
and re-identified the flies from T. califor- 
nica as T. arizonensis; moreover, Goeden et 
al. (1998b) further supported this identifi- 
cation based on the different tribal- and sep- 
arate, subtribal-level, host-plant affiliations 



VOLUME 101. NUMBER 1 



77 



of the three biotypes of T. actinobola in 
southern California, which exclude Trixis 
californica in the tribe Mutisieae (see be- 
low). 

Nine (10.6%) of 85 S, but none of 65 9 
of Trupanea arizonensis in the research col- 
lection of RDG reared from flower heads of 
Trixis californica have only one dark ray, 
not two rays, contrary to Foote et al. 
(1993), extending into cell dm from vein 
CuA|. Moreover, the single, remaining, 
what is otherwise called the distal ray 
(Foote et al. 1993), is broken in cell dm and 
does not reach vein CuA, in 39 6 (45.9%) 
and 18 $ (27.7%). There is no dark spot 
on vein CuA, or on the wing margin in line 
with the broken distal ray in an additional 
10 S (11.8%) and 3 ? (4.6%), or with such 
a dark spot on CuA, or on the wing margin 
in line with the broken distal ray in an ad- 
ditional 2 6 (2.4%) and 3 9 (4.6%), re- 
spectively. The proximal ray usually ex- 
tends, though sometimes very limitedly, 
into cell dm in 76 6 (89%) and all 65 9, 
and there is usually (not always, Foote et 
al. 1993) a dark spot on vein CuA, in line 
with it; however, in 3 9 (4.6%) the broken 
proximal ray is reduced to a spot in the 
middle of cell dm in line with a spot on 
vein CuA|, and another 2 6 (2.3%) and 1 
9 (1.5%) lack a dark spot in line with it. 
Therefore, all females and most males of T. 
arizonensis can still be distinguished from 
other Trupanea spp. by use of the separate 
keys for the two sexes in Foote et al. (1993) 
if one is aware of the above variations and 
perceives two dark rays, however abbrevi- 
ated, extending into cell dm, neither of 
which necessarily crosses vein CuA,, much 
less continues to the hind wing margin. 
Males with only one dark ray extending 
into cell dm from vein CuA,, like the nine 
noted above, will run to T. actinobola, as 
they did earlier for Goeden and Ricker 
(1989). Couplet 8 (pp. 421 and 424 in Foote 
et al. 1993) distinguishing T. maciiligera 
Foote from T. arizonensis apparently re- 
mains valid; although the angles formed by 
the proximal and distal rays, not the dis- 




Fig. 1. Egg ol Trupanea arizonensis: (A) habitus, 
pedicel to left; (B) pedicel. 

tances between them along vein CuA,, 
should be compared with specimens of T 
arizonensis having proximal or distal rays 
or both broken in cell dm. 

Immature stages. — The immature stages 
of T arizonensis heretofore have neither 
been described nor illustrated. 

Egg: Seventy eggs of T arizonensis dis- 
sected from heads of T californica were 
white, opaque, smooth; elongate-ellipsoi- 
dal, 0.70 ± 0.006 (range, 0.54-0.85) mm 
long, 0.15 ± 0.002 (range, 0.12-0.23) mm 
wide, smoothly rounded at tapered basal 
end (Fig. lA), pedicel 0.02 mm long, with 
single row of subcircular aeropyles (Fig. 
IB). 

The egg of T. arizonensis is narrow like 
the eggs of T actinobola (Goeden et al. 
1998b) and T pseiidovicina (Goeden and 
Teerink 1998), but is much longer than both 
of these species. The egg body in T. ari- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. First instar of Trupanea arizonensis: (A) habitus, anterior end to left: (B) gnathocephalon, anterior 
view, 1, dorsal sensory organ, 2, terminal sensory organ, 3, pit sensory organ, 4. lateral sensory organ, 5, stomal 
sense organ; (C) gnathocephalon, ventral view, 1, anterior sensory lobe, 2, mouth hook, 3, median oral lobe, 4, 
labial lobe, 5, pit sensillum; (D) caudal segment, 1, stelex sensillum, 2, rima, 3. interspiracular process, 4, 
intermediate sensory complex. 



zonensis tapers smoothly into the pedicel, 
more so than in the other Trupanea species 
previously studied (Goeden and Teerink 
1997, 1998; Goeden et al. 1998a, b; Knio 
et al. 1996a; Teerink and Goeden 1998). 
The pedicel is similar to these other con- 
geners in having a single row of aeropyles 
(Goeden and Teerink 1997, 1998; Goeden 
et al. 1998a, b; Teerink and Goeden 1998), 
with the exception of T. bisetosa (Coquil- 
lett), which has one or two rows (Knio et 
al. 1996a). 

First instar. White, elongate-cylindrical, 
rounded anteriorly and posteriorly (Fig. 
2A), minute acanthae circumscribe inter- 
segmental lines (Fig. 2A); gnathocephalon 
smooth (roughness in Fig. 2 is an artifact), 
lacking rugose pads (Fig. 2C); dorsal sen- 



sory organ a dome-shaped papilla (Fig. 2B- 
1); anterior sensory lobe (Fig. 2C-1) bears 
terminal sensory organ (Fig. 2B-2), pit sen- 
sory organ (Fig. 2B-3), lateral sensory or- 
gan (Fig. 2B-4), and supralateral sensory 
organ; stomal sense organ indistinct (Fig. 
2B-5); mouth hooks bidentate (Fig. 2C-2); 
median oral lobe laterally flattened (Fig. 
2C-3), labial lobe (Fig. 2C-4) attached to 
median oral lobe; pit sensillum laterad of 
anterior sensory lobe (Fig. 2C-5); prothorax 
lacking rugose pads, few minute acanthae 
ventrad of mouth lumen; stelex sensilla cir- 
cumscribe caudal segment in 2-dorsal, 4- 
ventral arrangement (Fig. 2D-1); posterior 
spiracular plates bear two ovoid rimae (Fig. 
2D-2), and four, spatulate interspiracular 
processes (Fig. 2D-3); intermediate sensory 



VOLUME 101, NUMBER 1 



79 



complex consists of a medusoid and stelex 
sensillum (Fig. 2D-4). 

The first instar of T. arizonensis is very 
similar to other congeners previously stud- 
ied in habitus and sensory structures (Goe- 
den and Teerink 1998; Goeden et al. 1998a, 
b; Knio et al. 1996a; Teerink and Goeden 
1998). The lateral spiracular complex was 
not seen. There are slight differences in the 
interspiracular processes. In T. jonesi, the 
interspiracular processes are greatly re- 
duced (Goeden et al. 1998a), in T. arizo- 
nensis and T. conjuncta the processes are 
single and spatulate (Teerink and Goeden 
1998), and in T. pseudovicina Hering, T. 
actinobola, T. bisetosa and T. nigricornis 
(Coquillett), the interspiracular processes 
are divided, each with 1-4 branches (Goe- 
den and Teerink 1997; Goeden et al. 1998b; 
Knio et al. 1996a). 

Third instar: White, elongate-cylindri- 
cal, tapering anteriorly, rounded posteriorly, 
minute acanthae circumscribe intersegmen- 
tal lines in bands increasing in width pos- 
teriorly (Fig. 3A); gnathocephalon conical, 
rugose pads dorsal and lateral to mouth lu- 
men (Fig. 3B-1); dorsal sensory organ a 
dome-shaped papilla (Fig. 3B-2, 3C-1); an- 
terior sensory lobe (Fig. 3B-3) bears the ter- 
minal sensory organ (Fig. 3C-2), pit sen- 
sory organ (Fig. 3C-3), lateral sensory 
organ (Fig. 3C-4), and supralateral sensory 
organ (Fig. 3C-5); stomal sense organ ven- 
trolaterad of anterior sensory lobe (Fig. 3B- 

4, 3C-6); mouth hooks tridentate (Fig. 3B- 

5, 3D-1); median oral lobe tapered anteri- 
orly (Fig. 3D-2); six pit sensilla circum- 
scribe gnathocephalon posterior to rugose 
pads (Fig. 3B-6); minute acanthae circum- 
scribe anterior margin of prothorax (Fig. 
3E-1); rugose pads (Fig. 3E-2) and a single 
row of verruciform sensilla (Fig. 3E-3) cir- 
cumscribe prothorax posteriorad of minute 
acanthae; additional verruciform sensilla on 
dorsal half of prothorax (Fig. 3E-4); ante- 
rior thoracic spiracle on posterior margin of 
prothorax bears three ovoid papillae (Fig. 
3E-5), meso- and metathorax circumscribed 
by verruciform sensilla; meso- and meta- 



thoracic lateral spiracular complexes consist 
of a spiracle (Fig. 3F-1), and three verru- 
ciform sensilla (Fig. 3F-2); abdominal lat- 
eral spiracular complex consists of a spira- 
cle (Fig. 3G-1) and two verruciform sensilla 
(Fig. 3G-2); caudal segment circumscribed 
by minute acanthae (Fig. 3H-1); posterior 
spiracular plates bear three ovoid rimae, ca. 
0.03 mm in length (Fig. 3H-2), and four 
interspiracular processes, each with 3-5 
branches, longest measuring 0.02 mm (Fig. 
3H-3); intermediate sensory complex con- 
sists of a stelex sensillum and a medusoid 
sensillum (not shown). 

The third instar of T. arizonensis is sim- 
ilar in general habitus to T. pseudovicina in 
being elongate-cylindrical, maybe even 
more attenuated (Goeden and Teerink 
1998). As with the third instars of each of 
the eight other Trupanea species previously 
studied, the lateral spiracular complex is 
unique to T. arizonensis, with three verru- 
ciform sensilla in the meso- and metathorax 
and two verruciform sensilla in the abdom- 
inal segments (Goeden and Teerink 1997, 
1998; Goeden et al. 1998a, b; Headrick and 
Goeden 1991; Knio et al. 1996a; Teerink 
and Goeden 1998). There are other differ- 
ences between individual species. Trupanea 
arizonensis differs from T actinobola in 
lacking elongated integumental petals; T. 
arizonensis has fewer rugose pads circum- 
scribing the prothorax, and the prothorax is 
completely circumscribed by minute acan- 
thae (Goeden et al. 1998b). Also, T. arizo- 
nensis differs from the other Trupanea spe- 
cies previously studied, in that the meso- 
and metathorax are circumscribed by ver- 
ruciform sensilla; whereas, only the meso- 
thorax in T. nigricornis was circumscribed 
by verruciform sensilla (Goeden and Teer- 
ink 1997, 1998; Goeden et al. 1998a, b; 
Headrick and Goeden 1991; Knio et al. 
1996a; Teerink and Goeden 1998). Trupa- 
nea arizonensis differs from its sympha- 
gous congener in Trixis californica, T. con- 
juncta, in not being finely punctate nor bar- 
rel-shaped, and T arizonensis also has 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Third iiistar of Trupanea arizcmensis: (A) habitus, anterior end to left; (B) gnathocephalon. antero- 
lateral view. 1 . rugose pads. 2. dorsal sensory organ. 3, anterior sensory lobe. 4, stomal sense organ, 5. mouth 
hook, 6. pit sensillum; (C) anterior sensory lobe, 1, dorsal sensory organ. 2, terminal sensory organ. 3. pit 
sensory organ. 4. lateral sensory organ. 5, supralateral sensory organ. 6, stomal sense organ; (D) gnathocephalon. 
ventral view, 1. mouth hook, 2, median oral lobe; (E) gnathocephalon, prothorax, lateral view. 1, minute acanthae. 



VOLUME 101, NUMBER 1 



81 



fewer papillae in the anterior thoracic spi- 
racle (Teerink and Goeden 1998). 

Pupariiim: Black, elongate-cylindrical, 
minute acanthae circumscribe intersegmen- 
tal lines, (Fig. 4A); anterior end bears the 
invagination scar (Fig. 4B-1), and anterior 
thoracic spiracles (Fig. 4B-2); caudal seg- 
ment circumscribed by minute acanthae 
(Fig. 4C-1); posterior spiracular plates bear 
three ovoid rimae (Fig. 4C-2) and four in- 
terspiracular processes, each with 3-5 
branches (Fig. 4C-3). Sixty-four puparia av- 
eraged 2.68 ± 0.02 (range, 2.28-3.24) mm 
in length; 0.92 ± 0.01 (range, 0.76-1.16) 
mm in width. 

Distribution and Hosts 

The distribution of T. arizonensis 
mapped in North America north of Mexico 
by Foote et al. (1993) is confined to several, 
mostly U.S. southern border locations in 
Arizona, California, and Texas. 

Goeden and Ricker (1989) reported T. 
arizonensis as T. actinobola from Trixis 
californica, and as mentioned above, Goe- 
den (1992) corrected this misidentification 
and thus provided the first and sole host- 
plant record for T. arizonensis. This host- 
plant belongs to the subtribe Nassauviinae 
in the tribe Mutisieae of the Asteraceae 
(Hickman 1993, Bremer 1994); this tribe 
rarely occurs in North America (Bremer 
1994), and is represented by only three spe- 
cies in California (Munz 1974), the second 
of which, Acourtia microcephala de- 
Candolle, does not host T. arizonensis 
(Goeden and Headrick 1991). The third 
plant species is the rare, unsampled, xero- 
phytic shrub, Hecastocleis shockleyi A. 
Gray (Hickman 1993, Bremer 1994). In 
comparison, T. actinobola, apparently is 
confined to three subtribes of the different 
tribe Astereae in California (Goeden and 



Teerink 1998b). Accordingly, T. arizonen- 
sis may be either a true monophage (one 
host-plant species) on Trixis californica, 
which occurs as far east as Texas in North 
America north of Mexico (Hickman 1993), 
or a near-monophage (one host-plant genus) 
on one or more of the 50 other congeners 
in the southwestern United States, Central, 
and South America and West Indies (Bre- 
mer 1994). Like several other tephritid spe- 
cies that we have studied, e.g., Trupanea 
conjimcta (Goeden 1987), T. pseiidovicina 
Hering (Goeden and Teerink 1998), and To- 
moplagia cressoni Aczel (Goeden and 
Headrick 1991), Trupanea arizonensis rep- 
resents a native southern California tephri- 
tid closely associated with a native host- 
plant, which is primarily distributed in 
Mexico and southward, where these tephri- 
tids remain little known. 

Biology 

Egg. — In 38 closed, preblossom, imma- 
ture flower heads, all 130 eggs were insert- 
ed pedicel-last between the tips of the phyl- 
laries, perpendicular to the receptacle, and 
among or within the florets and pappus 
(Fig. 5A). The diameters of the receptacles 
of 32 of these flower heads containing eggs 
averaged 2.6 ±0.1 (range, 1.3-3.7) mm, 
and the 38 infested heads contained an av- 
erage of 3.4 ± 0.3 (range, 1-9) eggs ovi- 
posited, mostly singly, or side-by-side in 
pairs (Figs. 5 A, B), but also in groups of 
up to five, by one or more females. 

Larva. — Upon eclosion, first instars tun- 
neled into a single floral tube or ovule of 
an immature floret (Fig. 5C). An average of 
2.1 ± 0.2 (range, 1-4) first instars was 
found feeding within 22 closed, preblossom 
flower heads. The receptacles of these 
heads averaged 3.1 ± 0.2 (range, 2.2-4.6) 
mm in diameter with an average of 18 ± 2 



2, rugose pads, 3. verruciform sensillum, 4, verruciform sensillum, 5, anterior thoracic spiracle; (F) metathorax, 
1, spiracle, 2, verruciform sensilla; (G) fifth abdominal segment, 1, spiracle, 2, verruciform sensilla; (H) caudal 
segment, 1, minute acanthae, 2, rima, 3, interspiracular process. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



V^n/' 



AccV Spot Magn WD 
lOOkV 3 38x 33,3 




Fig. 4. Puparium of Tnipanea cirizoiiensis: (A) 
habitus, anterior end to right; (B) anterior end, 1, in- 
vagination scar, 2, anterior thoracic spiracle; (C) caudal 
segment, K minute acanthae, 2, rima, 3, interspiracular 
process. 



(range, 15-24) florets, of which an average 
of only 1.6 ± 0.2 (range, 1-3) florets, or 
9% (range, 4-17%), were damaged. No re- 
ceptacles within these 14 infested flower 
heads was pitted by first-instar feeding. 



Second instars fed solitarily at the distal 
ends of individual ovules or soft achenes 
within different florets of separate preblos- 
som and open flower heads, respectively. 
Their mouthparts were directed towards the 
receptacles. Receptacles of 16 flower heads 
containing second instars were not pitted 
and averaged 3.4 ± 0.2 (range, 2.3-4.8) 
mm in diameter. These flower heads con- 
tained an average of 1.6 ± 0.2 (range, 1-3) 
second instars that had damaged an average 
of 1.8 ± 0.2 (range, 1-4) ovules, or 11% 
(range, 6-22%) of an average total of 17 ± 
0.8 (range, 12-20) ovules per flower head. 

Most third instars also confined their 
feeding to the same single, separate ovule 
or soft achene at the centers, and less com- 
monly at the margins, of preblossom or 
open flower heads (Fig. 5D). Third instars 
fed with their long axes oriented perpendic- 
ular to and mouthparts directed towards the 
receptacles, and on the distal parts of the 
ovules or soft achenes, well above the re- 
ceptacles (Fig. 5D). In 28 flower heads av- 
eraging 4.0 ± 0.1 (range, 3.1-5.0) mm in 
diameter and containing an average of 1.5 
± 0.2 (range, 1-5) third instars, an average 
of 1.6 ± 0.2 (range, 1-5) ovules were dam- 
aged, or 9% (range, 4-25%). Thus, each 
larva confined its feeding to the distal parts 
of a single floret, at most including corolla 
tube and contents, pappus, and upper part 
of the ovule (Fig. 5D); the ovule continued 
to grow and functioned as a nutrient sink 
(Harris and Shorthouse 1996) augmented 
by the larva which fed mainly on sap drawn 
to the cuplike depression in the distal end 
of the ovule (Fig. 5D). This mode of feed- 
ing is the least damaging to host-plant re- 
production, as well as the most exquisitely 
evolved manner of sap-feeding yet reported 
among florivorous Tephritidae (Headrick 
and Goeden 1998). This manner of larval 
feeding, first reported by Headrick and 
Goeden (1990b), apparently is facultative, 
undetected, or absent in some florivorous 
species, e.g., Paracantha gentilis Hering 
(Headrick and Goeden 1990a, b), Trupanea 
nigricomis and T. bisetosa (Knio et al. 



VOLUME 101, NUMBER 



83 




Fig. 5. Lite stages of Triipanea arizonensis in or on r//.v/.v califumica: (A) one pair of eggs (arrow) inserted 
among immature florets in closed, preblossom flower head; (B) two pairs of eggs inserted among florets in 
closed, preblossom flower head; (C) first instar tunneling in single floret (anow); (D) two third instars feeding 
on separate soft achenes; (E) single puparium atop stunted achene in flower head; (F) adult female at rest on 
flower head. Lines = 1 mm. 



(1996b); although it also has been reported 
in some gallicolous species, e.g., Aciuhna 
thoracica Curran (Headrick and Goeden 
1993), and is facilitated by specialized 
mouthparts in larval Tephritinae (Headrick 
and Goeden 1990a, 1993, 1998). 

Upon completing feeding, the larvae ori- 
ented their anterior ends away from the re- 
ceptacles, retracted their mouthparts, and 
pupariated (Fig. 5E). 

Pupa. — Flower heads containing puparia 
(Fig. 5E) contained the greatest amounts of 
damage produced by the seed-feeding lar- 



vae of T. arizonensis. The receptacles of 41 
open and postblossom flower heads con- 
taining puparia averaged 4.4 ± 0.1 (range, 
2.9-6.2) mm in diameter and bone an av- 
erage total of 19.2 ± 0.6 (12-26) soft 
achenes, of which an average of 2.2 ± 0.3 
(range, 1-9) soft achenes or 13% (range, 4- 
58%) were damaged. Again, no receptacles 
were fed upon. These heads contained an 
average of 1.8 ± 0.2 (range, 1-7) puparia. 
Most puparia of T. arizonensis were found 
near the centers of the flower heads, and all 
had their anterior ends facing away from 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



the receptacles, and their long axes were 
perpendicular to the receptacles (Fig. 5E). 

Adult. — Adults (Fig. 5F) emerged from 
mature, postblossom flower heads, and 
were long-lived under insectary conditions, 
as 15 unmated males lived an average of 89 
± 14 (range, 11-194) days, and six virgin 
females averaged 126 ± 7 (range, 1 10-150) 
days. Like several other, monophagous and 
nearly monophagous congeners studied 
(Goeden 1988, Goeden and Teerink 1997, 
1998), the longevities of these flies were 
among the longer averages and maxima for 
adults that we have recorded for native spe- 
cies of nonfrugivorous Tephritidae from 
southern California. Such longevities are 
commensurate with the aggregative type of 
life cycle ascribed below to this tephritid. 
The premating and mating behaviors of T. 
arizonensis were not studied in the field, 
and again, like most congeners that we have 
studied, adults would not mate in petri dish 
arenas otherwise so useful with many other, 
noncongeneric, nonfrugivorous species 
(Headrick and Goeden 1994). 

Seasonal history. — The life cycle of T. 
arizonensis in southern California appears 
to follow an aggregative pattern in which 
the long-lived adults in reproductive dia- 
pause over-summer in riparian habitats and 
mountain meadows. They return to lower 
elevations in the fall, and following the on- 
set of winter rainfall, aggregate on Trixis 
californica in the low-elevation Colorado 
Desert in winter (January-February) to 
mate and later to oviposit (March-April) 
(Headrick and Goeden 1994). A single an- 
nual generation is produced each year and 
most of the life span of T. arizonensis is 
spent as an adult. 

Natural enemies. — Pteromalus sp. (Hy- 
menoptera: Pteromalidae) was reared from 
individual puparia of T. arizonensis as a 
solitary, larval-pupal endoparasitoid; how- 
ever, at least some of the other species of 
parasitoids reported from Tomoplagia cres- 
soni Aczel (Goeden and Headrick 1991) 
and Trupanea conjuncta (Goeden 1987), 
which co-occur in symphagy in the heads 



of Trixis californica (Goeden and Ricker 
1989, Goeden 1997), probably also parasit- 
ize T. arizonensis. 

Acknowledgments 

We thank Andrew C. Sanders, Curator of 
the Herbarium, Department of Botany and 
Plant Sciences, University of California, 
Riverside, for identification of plants men- 
tioned in this paper. The parasitoids were 
identified by Harry E. Andersen, 
Huntington Beach, California. We also are 
grateful to Kristine Gilbert for technical as- 
sistance, and to D. H. Headrick for his help- 
ful comments on an earlier draft of this pa- 
pen 

Literature Cited 

Bremer. K. 1994. Asteraceae Cladistics & Classifica- 
tion. Timber Press, Inc. Portland. Oregon. 

Cavender, G. L. and R. D. Goeden. 1982. Life history 
of Trupanea hisetosa (Diptera: Tephritidae) on 
wild sunflower in southern California. Annals of 
the Entomological Society of America 75: 400- 
406. 

Foote, R. H. 1960. A revision of the genus Trupanea 
in America north of Mexico. United States De- 
partment of Agriculture Technical Bulletin 1214. 
29 pp. 

Foote, R. H. and F L. Blanc. 1963. The fruit flies or 
Tephritidae of California. Bulletin of the Califor- 
nia Insect Survey 7, 117 pp. 

Foote, R. H., F L. Blanc, and A. L. Norrbom. 1993. 
Handbook of the Fruit Flies (Diptera: Tephritidae) 
of America North of Mexico. Cornell University 
Press, Ithaca, New York. 

Goeden, R. D. 1985. Host-plant relations of Trupanea 
spp. (Diptera: Tephritidae) in southern California. 
Proceedings of the Entomological Society of 
Washington 87: 564-571. 

. 1987. Life history of Trupanea conjuncta 

(Adams) on Trixis californica Kellogg in southern 
California (Diptera: Tephritidae). Pan-Pacific En- 
tomologist 63: 284-291. 

. 1988. Life history of Trupanea impetfecta 

(Coquillett) on Bebbia juncea (Bentham) Greene 
in the Colorado Desert of southern California 
(Diptera: Tephritidae). Pan-Pacific Entomologist 
64: 345-351. 

. 1992. Analysis of known and new host re- 
cords for Trupanea from California (Diptera: Te- 
phritidae). Proceedings of the Entomological So- 
ciety of Washington 94: 107-1 18. 

. 1997. Symphagy among florivorous fruit flies 

(Diptera: Tephritidae) in southern California, pp. 



VOLUME 101, NUMBER 1 



85 



39-52 //; Dettner, K., G. Bauer, and W. Vokl, eds.. 
Vertical Food Web Interactions: Evolutionary Pat- 
terns and Driving Forces. Ecological Studies 130, 
Springer- Verlag, Heidelberg, Germany. 

Goeden. R. D. and D. H. Headrick. 1991. Notes on 
the biology, hosts, and immature stages of To- 
moplagia cressoni (Aczel) in southern California 
(Diptera: Tephritidae). Proceedings of the Ento- 
mological Society of Washington 73: 549-558. 

Goeden, R. D. and D. W. Ricker. 1989. Phytophagous 
insect faunas of the desert shrubs, Bebbia juncea 
and Trixis californica in southern California. An- 
nals of the Entomological Society of America 82: 
325-321. 

Goeden, R. D. and J. A. Teerink. 1997. Life history 
and description of immature stages of Tntpanea 
signata Foote (Diptera: Tephritidae) on Gnaphal- 
iiim luteo-album L. in southern California. Pro- 
ceedings of the Entomological Society of Wash- 
ington 99: 748-755. 

. 1998. Life history and description of imma- 
ture stages of Trupanea pseiidovicina Bering 
(Diptera: Tephritidae) on Porophyllum gracile 
Bentham (Asteraceae) in southern California. Pro- 
ceedings of the Entomological Society of Wash- 
ington 100: 361-372. 

Goeden, R. D., J. A. Teerink, and D. H. Headrick. 
1998a. Life history and description of immature 
stages of Trupanea jonesi Curran (Diptera: Te- 
phritidae) on native Asteraceae in southern Cali- 
fornia. Proceedings of the Entomological Society 
of Washington 100: 126-140. 

. 1998b. Life history and description of im- 
mature stages of Trupanea actinobola (Loew) 
(Diptera: Tephritidae) on Acamptopappus sphaer- 
ocephalus (Harvey and Gray) Gray (Asteraceae) 
in southern California. Proceedings of the Ento- 
mological Society of Washington 100: 674-688. 

Harris, P. and J. D. Shorthouse. 1996. Effectiveness of 
gall inducers in weed biological control. Canadian 
Entomologist 128: 1021-1055. 

Headrick, D. H. and R. D. Goeden. 1990a. Description 
of the immature stages of Paracantha gentilis 
(Diptera: Tephritidae). Annals of the Entmological 
Society of America 83: 220-229. 



. 1990b. Resource utilization by larvae of Par- 
acantha gentilis (Diptera: Tephritidae) in capitula 
of Cirsium californicum and C. proteanum (As- 
teraceae) in southern California. Proceedings of 
the Entomological Society of Washington 92: 
512-520. 

. 1991. Life history of Trupanea californica 

Malloch (Diptera: Tephritidae) on Gnapluilium 
spp. in southern California. Proceedings of the En- 
tomological Society of Washington 93: 559-570. 

. 1993. Life history and description of imma- 
ture stages of Aciurina thoracica Curran (Diptera: 
Tephritidae) on Baccharis sarothroides Gray in 
southern California. Proceedings of the Entomo- 
logical Society of Washington 95: 210-222. 

. 1994. Reproductive behavior of California 

fruit flies and the classification and evolution of 
Tephritidae (Diptera) mating systems. Studia Dip- 
terologica 1(2): 194-252. 

. 1998. The biology of nonfrugivorous tephritid 



fruit flies. Annual Review of Entomology 43: 
217-241. 

Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- 
versity of California Press. Berkeley and Los An- 
geles. 

Knio, K. M., R. D. Goeden, and D. H. Headrick. 
1996a. Descriptions of immature stages of Tru- 
panea nigricornis and T. bisetosa (Diptera: Te- 
phritidae) from southern California. Annals of the 
Entomological Society of America 89: 1 — 11. 

. 1996b. Comparative biologies of the cryptic, 

sympatric species, Trupanea nigricornis and T. 
bisetosa (Diptera: Tephritidae) in southern Cali- 
fornia. Annals of the Entomological Society of 
America 89: 252-260. 

Malloch, J. R. 1942. Notes on two genera of American 
flies of the family Trypetidae. Proceedings of the 
United States National Museum 92(3123): 1-20. 

Munz, P. A. 1974. A Flora of Southern California. Uni- 
versity of California Press, Berkeley and Los An- 
geles. 

Teerink, J. A. and R. D. Goeden. 1998. Description of 
immature stages of Trupanea conjuncta Adams 
(Diptera: Tephritidae). Proceedings of the Ento- 
mological Society of Washington 100: 431-438. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999. pp. 86-93 

TWO NEW SPECIES OF AGARODES BANKS 
(TRICHOPTERA: SERICOSTOMATIDAE) FROM SOUTHEASTERN 

UNITED STATES 

A. C. Keth and S. C. Harris 

Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. (e-mail: 
harris@mail.clarion.edu) 



Abstract. — Two new species, Agarodes logani, from the eastern panhandle of Florida, 
and Agarodes tuskaloosa, from west-central Alabama, are described, illustrated, and com- 
pared to congeners. The genus now totals twelve species, all restricted to eastern North 
America. 

Key Words: Trichoptera, Sericostomatidae, Agarodes, new species, southeast 



The genus Agarodes Banks is restricted 
to eastern North America and ranges from 
southern Canada to the southern United 
States (Ross and Scott 1974). Agarodes lar- 
vae prefer smaller, spring-fed streams with 
a medium current and sandy substrate but 
they have been occasionally collected in 
larger streams fed by surface water and in 
the sandy, depositional areas of lakes. 

Ten species of Agarodes were previously 
recognized (Harris 1987) with all but A. 
distincta Ulmer and A. grisea Banks re- 
stricted to southeastern United States (Ross 
and Scott 1974). In Alabama, five species 
occur, A. crassicornis Walker, A. libalis 
Ross and Scott, A. alabamensis Harris on 
the Coastal Plain; A. grisea Banks in north- 
ern Alabama; and A. stannardi Ross in a 
restricted, northwestern region of the state. 
To this list from Alabama an apparently 
rare, new species is added, A. tuskaloosa. 
This new species is currently known only 
from the type locality, on a small, spring- 
fed stream at the northern edge of the 
Coastal Plain. In Florida, three species oc- 
cur, A. crassicornis Walker and A. libalis 
Ross and Scott at scattered localities; and 
A. ziczac Ross and Scott known only from 



the type locality in the central panhandle 
region. To this Florida list, A. logani. a new 
species, is added. This species is currently 
known only from the type locality, a small 
spring-run in a deep ravine, in the eastern 
panhandle region. Agarodes tuskaloosa and 
Agarodes logani are members of the sub- 
genus Agarodes Banks (Ross and Wallace 
1974) based on the small antennal scape 
and slender mesal lobe of the maxillary 
palp. Both were collected with a black light. 
Type material is deposited at the National 
Museum of Natural History, Smithsonian 
Institution, Washington D.C. Terminology 
follows that of Schmid (1980) and Ross and 
Scott (1974). 

Agarodes logani Keth and Harris, 
new species 

(Figs. 1, 4) 

Diagnosis. — This species resembles A. 
stannardi but differs, primarily, in the elon- 
gate, strongly curved dorsomesal process of 
the mesal processes of the inferior append- 
age. The dorsomesal process (Fig. 1) of A. 
logani is about twice the length seen in A. 
stannardi (Fig. 2) and much more serrate 
ventrally. The female is very similar to A. 



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Fig. 1. Agarodes logani, male genitalia, a) lateral, b) ventral, c) dorsal, d) phallus. 



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Fig. 2. Agarodes stannardi. male genitalia, a) lateral, b) ventral, c) dorsal, d) phallus. 



VOLUME 101. NUMBER 1 



89 



stannardi differing only in the pair of short 
processes extending from apex of the dor- 
sum of segment IX. These processes in A. 
stannardi are divergent and basally lobate 
in dorsal view (Fig. 3) and in A. logani nar- 
row and lacking basal lobes (Fig. 4). 

Male. — Length 9-1 1 mm. Body, legs, 
and head golden to dark brown. Antenna 
brown with 43 segments, scape small and 
rectangular. Labial palpus 0.9 mm long; 
maxillary palpus 0.5 mm in length, oblong, 
and bearing 2 small inner processes. Wings 
golden to dark brown; forewing with exten- 
sive peg-like setae in anal region and along 
M and Cu veins. Abdominal segment VIII 
annular. Segment IX inserted within seg- 
ments VII and VIII, reduced, and dorsally 
fused with base of segment X. Segment X 
elongate and narrow in dorsal view, divided 
apically; in lateral view sinuate dorsally and 
lightly sclerotized, tapering to a point and 
curving slightly dorsad at apex. Genitalia as 
in Fig. 1 with preanal appendage narrow 
and elongate extending nearly half length 
of tergum X, curving dorsad. Inferior ap- 
pendage uniform over entire length in lat- 
eral view; in ventral view united basally, 
slender anteriorly, widening midway, and 
narrowing posteriorly. Mesal processes bi- 
partite; dorsomesal process over % length 
of inferior appendage, slender and narrow- 
ing to acute apex with large, serrate, ven- 
trad projections from % of length to apex 
and curving markedly ventrad in lateral 
view. Ventromesal process thin and nearly 
equal in length to dorsomesal process, di- 
vergent, curving markedly mesad at tip, 
slightly serrate distally at midlength and 
dorsally at apex. Phallus long and cylindri- 
cal, widening at membranous apex. 

Female. — Length, color, and general 
structure as in male. Genitalia as in Fig. 4 
with cerci long and contiguous, mesally 
merging with a short lobe. Apex of tergum 
IX having a pair of short processes pro- 
jecting posteriorly from dorsum; processes 
narrow and fused basally, lacking basola- 
teral lobes. Genital chamber narrow with 
margins folded, sclerotized, and curving 



mesad in ventral view, a pair of sclerous, 
scoop-like lobes diverging from base in 
dorsal view. 

Immatures. — Unknown 

Etymology. — Latin: of Logan, primary 
author's son 

Holotype. — d, Florida, Gadsden County, 
headwaters of Quincy Creek, 7 km. north 
Quincy at Florida A&M Research and Ex- 
tension Center, N30°39'27", W84°36'50", 
19 April 1994, Pescador and Rasmussen. 

Paratypes. — Florida, same as above, 1 S ; 
same, but N30°39'19", W84°36'51", 6 Oc- 
tober 1993, Jones, Pecador, and Rasmussen, 
1 $. 

Distribution. — Agarodes logani is known 
only from the type locality. More speci- 
mens have been found in the same area in- 
cluding Agarodes larvae which, upon rear- 
ing, may be found to be A. logani. 

Agarodes tuskaloosa Keth and Harris, 
new species 

(Fig. 5) 

Diagnosis. — This species resembles A. 
stannardi and the preceding new species, 
but it differs from A. stannardi in the much 
thinner inner process of the mesal process 
of the inferior appendage and from A. lo- 
gani in the presence of a short basomesal 
process (Fig. 5). The dorsal and ventrome- 
sal processes of A. tuskaloosa are like those 
of A. stannardi but twice the length. 

Male. — Length 11-13 mm. Body, legs 
and head golden to reddish brown. Antenna 
dark brown with 43 segments, scape small 
and trapezoidal. Labial palpus 1.0 mm long; 
maxillary palpus 0.6 mm in length, oblong. 
Wings light brown. Abdominal segment 
VIII annular. Segment IX inserted within 
segments VII and VIII, reduced, and dor- 
sally fused with base of segment X. Seg- 
ment X elongate and narrow in dorsal view, 
split apically; narrowing to elongate, acute 
apex in lateral view. Genitalia as in Fig. 5 
with preanal appendage narrow, elongate, VS 
length of tergum X, uniform over entire 
length in lateral view. Inferior appendage 
narrow basally, broadening toward apex in 



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Fig. 3. Agarodes stanuanU. female genitalia, a) lateral, b) ventral, c) dorsal. 



VOLUME 101. NUMBER I 



91 




Fig. 4. Agarodes logani, female genitalia, a) lateral, b) ventral, c) dorsal. 



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Fig. 5. Agarodes tuskaloosa, male genitalia, a) lateral, b) ventral, c) dorsal, d) phallus. 



VOLUME 101. NUMBER 1 



93 



lateral view; in ventral view slender and 
united basally, broadening % of length, and 
tapering to acute apex. Mesal processes tri- 
partite; basomesal process short and acute 
with slightly serrate apex dorsally, dorso- 
mesal process % length of inferior append- 
age and strongly curved in lateral view, 
apex slightly serrate ventrally; inner process 
thin, uniform, and elongate. Phallus long 
and cylindrical, widening slightly at mem- 
branous apex. 

Female. — Unknown 

Immatures. — Unknown 

Etymology. — Native American spelling: 
of Tuscaloosa region 

Holotype. — S, Alabama, Tuscaloosa 
County, Big Sandy Creek, 7.2 km. south of 
Coaling, on unmarked county road, 15 May 
1991, Harris. 

Paratype. — Alabama, locality and date as 
holotype, 1 6. 

Distribution. — Agarodes tuskaloosa is 
known only from the type locality. Exten- 
sive collecting at the type locality and else- 
where in Big Sandy Creek has yielded no 
additional specimens. 



Acknowledgments 
The Department of Biology of Clarion 
University of Pennsylvania provided equip- 
ment and facilities for this study and is 
thankfully recognized. Specimens for com- 
parison were provided by Dr. Brian Armi- 
tage at the Ohio Biological Survey and by 
Dr. John Morse at Clemson University. 
Thanks also to Jerome Jones, Manuel Pes- 
cador, and Andrew Rasmussen, of Florida 
A & M University, for specimen collections 
in northern Florida. 

Literature Cited 

Harris, S. C. 1987. A new species of Agarodes (Tri- 
choptera: Sericostomatidae) from southeastern 
United States. Proceedings of the Entomological 
Society of Washington 89( 1 ): 74-76. 

Ross, H. H. and D. C. Scott. 1974. A review of the 
caddisfly genus Agarodes with descriptions of 
new species (Trichoptera: Sericostomatidae). Jour- 
nal of the Georgia Entomological Society 9: 147- 
155. 

Ross, H. H. and J. B. Wallace. 1974. The North Amer- 
ican genera of the family Sericostomatidae (Tri- 
choptera). Journal of the Georgia Entomological 
Society 9: 42-48. 

Schmid, F. 1980. Genera des Trichopteres du Canada 
et des etats adjacents, pt. 7: 296 pp. //; les insectes 
et arachnides du Canada. Agriculture Canada, Ot- 
tawa. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999, pp. 94-98 

THE HIGHER CLASSIFICATION OF THE ALYDIDAE 
(HEMIPTERA: HETEROPTERA) 

Carl W. Schaefer 

Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, 
CT 06269-3043, U.S.A. (e-mail: schaefer@uconnvm.uconn.edu) 



Abstract. — Characters are taken from the literature and their derived states determined. 
The following higher classification of the Alydidae best agrees with the data: family 
Alydidae; subfamilies Alydinae and Micrelytrinae, the Alydinae with tribes Alydini and 
Daclerini; the Micrelytrinae with tribes Micrelytrini and Leptocorisini; the latter with 
subtribes Leptocorisidi and Noliphidi. 

Key Words: Alydidae, higher classification, Leptocorisini, Micrelytrinae, Noliphini, Da- 
clerini 



Until 1965 the family Alydidae had been 
variously treated as a subfamily of Corei- 
dae, or as a family in its own right. How- 
ever, when treated as a family the habit of 
subfamilial treatment persisted, and the 
lower categories were considered tribes, not 
subfamilies (see Schaefer [1965] for the 
taxonomic history), hi 1965 I presented ev- 
idence for family rank of the group, and 
recognized two subfamilies, Alydinae and 
Micrelytrinae, the latter with two tribes, Mi- 
crelytrini and Leptocorisini. 

In the same year, Ahmad (1965) pub- 
lished his revision of Leptocorisa and its 
relatives. He treated this group as a subfam- 
ily, of status equal to Alydinae isensii mihi) 
and Micrelytrinae (Micrelytrini met). He 
discussed these three subfamilies very brief- 
ly, but did not give arguments for treating 
them as of equal rank. Ahmad and I had 
not seen each other's papers, and therefore 
neither of us could consider the higher-rank 
treatment of the other. 

Since 1965, Ahmad's view has prevailed, 
with a lapse in 1979, when Ahmad et al. 
(1979) treated Leptocorisini and Micrelytri- 
ni as tribes in the subfamily Leptocorisinae. 



Nevertheless, most post- 1965 authors treat- 
ed the three groups as subfamilies, perhaps 
because Ahmad's (1965) study of the Lep- 
tocorisinae (sensu suo) was more detailed 
than mine of the entire Alydidae. Indeed, I 
myself subsequently treated the three 
groups as subfamilies (see, for example, 
Schaefer 1972, 1980, Schaefer et al. 1989, 
but see Schaefer 1996); most notably, the 
three subfamilies are recognized in the cat- 
alog of Nearctic Heteroptera (Froeschner 
1988). (Note: the change by Henry and 
Froeschner [1992] from Leptocorisinae to 
Leptocorinae is surely incorrect: the generic 
name is Leptocorisa, not Leptocoris [which 
is a genus in Rhopalidae].) 

In 1993, Li and Zheng published a study 
of alydid phylogeny. In it they concluded 
"that Schaefer's (1965) division of this 
family into two subfamilies — Alydinae and 
Micrelytrinae — is reasonable." Their 
"cladograms do not support Ahmad's 
(1965) and some other authors' contention 
that the Alydidae be divided into three sub- 
families — Alydinae, Leptocorisinae, and 
Micrelytrinae" (quoted from the English 
abstract of Li and Zheng [1993]). Since the 



VOLUME 101. NUMBER 1 



95 



appearance of this paper. Prof. Zheng has 
most generously provided me with an En- 
ghsh translation of it. 

Li and Zheng (1993) discuss and illus- 
trate many characters in some detail, thus 
adding substantially to the already rich lit- 
erature on Alydidae (see references in Table 
1). Accordingly, I have extracted characters 
from this literature and have attempted to 
determine their derived states, in an attempt 
to establish better the Alydidae's higher 
classification. 

Methods 

I extracted from the literature on the Al- 
ydidae characters whose states could be 
tabulated and whose polarities for the most 
part could be determined. All these char- 
acters are ones important in the higher sys- 
tematics of Coreoidea and, indeed, of Pen- 
tatomomorpha (the heteropteran infraorder 
to which Coreoidea belongs; see Henry 
1997). Sample size is of course a problem; 
all genera and many species of Alydinae are 
described in Schaffner's dissertation (1964), 
but only for the Leptocorisini are all mem- 
bers of the group treated and their mor- 
phology described, in Ahmad's (1965) 
monograph. In particular, the Micrelytrini 
need revision, with attention given to mor- 
phological features of systematic impor- 
tance in other alydids. 

In polarizing the character states I take 
the Coreidae as the outgroup. Henry (1997) 
presents convincing evidence that this fam- 
ily is the sibling group of Alydidae. Other 
arguments for my polarizing occur in the 
references and in footnotes to Table 1. 

Results and Discussion 

Twenty-eight characters and their states 
are in Table 1 (where I anticipate my con- 
clusions by treating Micrelytrini and Lep- 
tocorisini as tribes and Alydinae as a sub- 
family). Of these characters, I polarized 23; 
three of the remainder I could not polarize 
(Distribution, Host plants. Rostral seg- 
ments); one (Scent gland auricle) is ambig- 
uous; and it is not clear to me if bifid (Mi- 



crelytrini) and trifid (Leptocorisini) medial 
projections of the genital capsule are sepa- 
rate advances over the (primitive) alydine 
condition, or whether the trifid condition is 
a further advance over the bifid (in which 
case, this character would group Micrely- 
trini and Leptocorisini together). 

Thirteen of the characters are autapo- 
morphies of one of the three groups. These 
autapomorphies are not the only ones defin- 
ing these groups, of course, because I was 
not seeking autapomorphies in the litera- 
ture. Nevertheless, the fact that Alydinae 
has eight autapomorphies, and the other two 
groups have fewer (Micrelytrini: 4; Lepto- 
corisini: 1), supports the subfamily status of 
Alydinae. 

Within the Alydinae, Ahmad et al. (1979) 
created a tribe, Daclerini, for the published 
genus Daclera and for another, unpublished 
genus. Although they present the new tribe 
as "MS," it is briefly described in their key 
and therefore appears to be valid. I have not 
seen specimens either of Daclera or of the 
undescribed genus. However, Li and Zheng 
(1993) write that Daclera has many apo- 
morphies not shared with other Alydinae. 
Therefore I treat Daclerini as a tribe in Al- 
ydinae, pending further study. 

Within the subfamily Micrelytrinae, Mi- 
crelytrini and Leptocorisini share six apo- 
morphies (seven, if the median projection 
condition is synapomorphic; see above), 
more than either group shares with Alydi- 
nae (Micrelytrini and Alydinae: 2; Lepto- 
corisini and Alydinae: 2). Three of these 
four synapomorphies shared by Alydinae 
and either Micrelytrini or Leptocorisini are 
head characters. The states of these char- 
acters vary considerably in the Coreoidea 
(Schaefer 1965), and their common posses- 
sion in Alydinae and one of the other tribes 
may therefore be homoplasious. If one ac- 
cepts that eight autapomorphies is a reason- 
able argument for subfamily status for Al- 
ydinae, then six synapomorphies should 
support subfamily status for Micrelytrini 
plus Leptocorisini. Moreover, four of these 
six synapomorphies are characters of the 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Table \. Differences and similarities among Micrelytrini, Leptocorisini. and Alydinae. Apomorphic states in 
boldface. 



Leptocorisini 



Alydinae 



Distribution 


tropical, subtropical 


tropical, subtropical 


Host plants 


Graminae (?) 


Graminae 


Head 






Midcephalic sulcus 


deep 


deep 


Head constricted 






basally 


yes 


rarely 


Head with "col- 






lar" 


no 


yes 


Ocelli on tubercle 


yes 


rarely 


Paraclypei well de- 


sometimes 


yes 


veloped 






Rostral segments 


2 > 3 -H 4; 4 = 
twice 3 


2<3 + 4;4 = 3 



tropical, subtropical; Schaffner 1964, Ah- 

some temperate mad 1965 

Leguminosae Schaefer 1979 

shallow or absent Li and Zheng 1993 



yes 

yes 
yes 

no 



< 3 4- 4; 4 > 3, 
rarely twice 3 



Li and Zheng 1993 

Schaefer 1965 

Li and Zheng 1993 

Li and Zheng 1993, 

Ahmad 1965 
Ahmad 1965, Schae- 
fer, unpubl. 



Thorax 
Hind femur 



Hind tibia 



not armed 
straight, untoothed 



not armed 
straight, untoothed 



Forewing media coriaceous basally coriaceous basally 
In membrane in membrane 

Forewing costa not fused to radius not fused to radius 

and media and media 

Scent gland peri- variable (no: Schae- no 

treme: lateral and fer 1965; yes: Li 

anterior auricles and Zheng 1993) 

fused 



spined or with stiff Schaffner 1964, Ah- 
setaC mad 1965 

usually curved and Schaffner 1964, Li 
with ventral and Zheng 1993 

tooth^ 

not coriaceous Li and Zheng 1993 



Abdomen 
Trichobothria (5th in a triangle 
sternum) 

Male genital capsule"^ 
Ventral rim with spine 

External opening posterior 



Dorsal wall 



sclerotized 



Median projection bifid 



in a triangle 



without spinc^^ 
dorsal 

sclerotized 



trifid 



fused'' 

yes 



in a line' 



Schaefer 1965, Li 
and Zheng 1993 

Schaefer 1965. Li 
and Zheng 1993 



Schaefer 1965. 1975 



without spine Schaefer 1980 

dorsal or postero- Schaefer 1980 
dorsal 



membranous 



single 



Cuplike sclerite 
with lateral pro- 
jections 

Segment 10 

Paramere 



no no 

sclerotized dorsally sclerotized dorsally 

apex not tuberculate apex not tuberculate 



Schaefer 1980. 

Schaefer et al. 

1989 
Schaefer 1980, 

Schaefer et al. 

1989 



yes Schaefer et al. 1989 

membranous dorsal- Schaefer 1980 

ly 

apex tuberculate Ahmad 1965, Li and 
Zheng 1993 



VOLUME 101, NUMBER 1 



97 



Table 1. Continued. 





Micrelytnni 


LcptOLonsini 


Al>dinac 




Rclctcncc 




Male aedeagus 














Vesica 


not slender, coiled' 


not slender, coiled' 


slender, straight' 


Schaefer 1965. 


Ah- 












mad 1965, L 


i and 












Zheng 1993 




Conjunctiva 


laterally with pair 


laterally with pair 


without these 


ap- 


Schaefer 1965, 


Ah- 




of asymmetrical 


of asymmetrical 


pendages 




mad 1965, L 


i and 




appendages 


appendages 






Zheng 1993 




Phallosoma 


dorsally with pair 


dorsally with pair 


without these 


ap- 


Schaefer 1965, 


Ah- 




of apically di- 


of apically di- 


pendages 




mad 1965, L 


i and 




rected append- 


rected append- 






Zheng 1993 






ages 


ages 










Female genitalia 














9th paratergite 


divided 


not divided 


not divided 




Schaefer 1965 




2nd valvula 


partly membranous 


sclerotized 


sclerotized 




Schaefer 1965 




Ring sclerites 


1 pair 


2 pairs 


1 pair 




Schaefer 1965 




Ring-sclerite sacs 


1 median sac 


1 pairs 


1 pair 




Schaefer 1965 





■' Curved and armed femora and tibia are uncommon in Coreoidea (except some Meropachydinae and some 
male Coreinae). 

^ Fusion of wing veins appears ipso facto to be more advanced than nonfusion. 

''The posterior abdominal trichobothria in Coreoidea are usually in a triangle (except Rhopalidae) (Schaefer 
1975). 

"• These features vary independently of one another (Schaefer 1980), and therefore may be treated as sepa- 
rate characters. 

''Spine is present in at least one leptocorisine (discussion in Schaefer 1980, p. 126). 

'According to Li and Zheng (1993), the apomorphic state of Alydinae differs from the apomorphic state of 
Micrelytrinae. 

s Ahmad (1965) writes that Leptocorisini have 0^ pairs of "'intervalvular sclerites"; I believe these are not 
the same as the sacs associated with the ring sclerites. 



male genitalia, a character complex always 
useful in heteropteran higher classification; 
common possession here is unlikely to be 
homoplasious In addition, other characters 
support the uniting of these two groups as 
a subfamily; these characters and the argu- 
ments based upon them, are not easily tab- 
ulated: see Schaefer (1965). 

Li and Zheng (1993) comment upon the 
antlike fascies of Alydinae, treating it as an 
autapomorphy of the group. It is true that 
immature alydines are antlike, but so are the 
adults of several Micrelytrini. In fact, there 
are two groups of Micrelytrini, one of 
somewhat or quite elongate insects, and the 
other of smaller often antlike insects 
(Schaefer 1996). Members of both groups 
occur in both the New and Old World trop- 
ics. By chance, Li and Zheng (1993) took 
as their representatives of Micrelytrini Mar- 



cius and Paramarcius, both members of the 
somewhat elongate (and nonantlike) group. 
Of great interest would be a study of the 
phylogenetic relationships among these 
four groups (New World and Old World 
antlike and elongate micrelytrines), be- 
tween the antlike micrelytrines and the Al- 
ydinae (whose nymphs are antlike), and be- 
tween the elongate micrelytrines and the 
Leptocorisini (most of whose genera are 
Old World tropical; the Noliphidi are some- 
what elongate and the Leptocorisidi are 
very elongate). I discussed some of these 
relationships earlier (Schaefer 1972). 

Classification 

As a result of this work, I suggest the 
following classification, in which the tribes 
of Ahmad's (1965) Leptocorisinae are re- 
duced to subtribes. 



98 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Family Alydidae Amyot et Serville 1 843 
Subfamily Alydinae Amyot et Serville 
1843 
Tribe Alydini 
Tribe Daclerini 
Subfamily Micrelytrinae Stal 1867 
Tribe Micrelytrini Stal 1867 
Tribe Leptocorisini Stal 1870 
Subtribe Leptocorisidi Stal 1870 
Subtribe Noliphidi Ahmad 1965 

Note: Based on their cladistic analysis, 
Li and Zheng (1993) suggest that Acestra, 
a genus placed uneasily in the Micrelytrini 
(see discussion in Li and Zheng [1993]), be 
removed from Micrelytrini and raised to 
tribal rank in the Micrelytrinae; they do not 
do this formally. Also, as I mentioned 
above, Li and Zheng (1993) found autapo- 
morphies in Daclera, the only described ge- 
nus now in Daclerini. These two genera 
should be studied more closely, as should 
the Micrelytrini as a whole. 

Acknowledgments 

I am deeply grateful to Zheng Le-yi for 
translating the Li and Zheng (1993) paper 
for me. Without his kindness and generos- 
ity, my paper had been difficult to under- 
take and impossible to complete satisfac- 
torily. I thank also Ann Harlan for word- 
processing this paper and for being a fa- 
vorite daughter. 

References Cited 

Ahmad, I. 1965. The Leptocorisinae (Heteroptera: Al- 
ydidae) of the world. Bulletin of the British Mu- 
seum (Natural History) Entomology Supplement 
5: 1-156. 

Ahmad. I., N. Abbas. M. U. Shadab, and A. A. Khan. 
1979. Generic and supergeneric keys with refer- 
ence to a checklist of alydid and stenocephalid 
fauna of Pakistan, Azad Kashmir and Bangladesh 
(Heteroptera: Coreoidea) with notes on their dis- 
tribution and food plants. Supplement of the En- 
tomological Society of Karachi, Pakistan 4(2): 1- 
19. 

Amyot, C. J. B. and A. Serville. 1843. Historic natu- 
relle des Insectes — Hemipteres. Librairies Ency- 
clopedique de Roret, Paris. 

Froeschner, R. C. 1988. Family Alydidae Amyot and 



Serville, 1843. The broad-headed bugs, pp. 4-11. 
//; Henry, T.J. and R. C. Froeschner, eds.. Catalog 
of the Heteroptera, or True Bugs, of Canada and 
the Continental United States. E.J. Brill, Leiden, 
The Netherlands. 

Henry, T. J. 1997. Phylogenetic analysis of family 
groups within the infraorder Pentatomomorpha 
(Hemiptera: Heteroptera), with emphasis on the 
Lygaeoidea. Annals of the Entomological Society 
of America 90: 275-301. 

Henry, T. J. and R. C. Froeschner. 1992. Corrections 
and additions to the "Catalog of the Heteroptera, 
or true bugs, of Canada and the continental United 
States." Proceedings of the Entomological Society 
of Washington 94: 263-272. 

Li, Xin-Zheng and Zheng Le-yi. 1993. Preliminary 
study on the phylogeny of Alydidae. Acta Zoo- 
taxonomia Sinica 18: 330-343 (Chinese, English 
summary.) [Note: I have used an English trans- 
lation prepared by Zheng Le-yi.] 

Schaefer, C. W. 1965. The morphology and higher 
classification of the Coreoidea (Hemiptera-Heter- 
optera). Part III. The families Rhopalidae, Alydi- 
dae, and Coreidae. Miscellaneous Publications of 
the Entomological Society of America 5( 1 ): 1-76. 

. 1972. Clades and grades in the Alydidae. 

Journal of the Kansas Entomological Society 45: 
135-141. 

. 1975. Heteropteran trichobothria (Hemiptera: 

Heteroptera). International Journal of Insect Mor- 
phology and Embryology 4: 193-264. 

. 1979. The host plants of the Alydinae, with a 

note on heteroptypic feeding aggregations (He- 
miptera: Coreoidea: Alydidae). Journal of the 
Kansas Entomological Society 53: 115-122. 

. 1980. The genital capsule of the Alydidae 

(Hemiptera: Heteroptera: Coreoidea). Internation- 
al Journal of Insect Morphology and Embryology 
9: 107-128. 

. 1996. A new species of Cydamiis. with a key 



to the species of the genus (Hemiptera: Alydidae). 
Annals of the Entomological Society of America 
89: 37-40. 

Schaefer, C. W., J. C. Schaffner, and I. Ahmad. 1989. 
The i4/vt/i/.v-group, with notes on the alydine gen- 
ital capsule (Hemiptera: Heteroptera: Alydidae). 
Annals of the Entomological Society of America 
82: 500-507. 

Schaffner, J. C. 1964. A taxonomic revision of certain 
genera of the tribe Alydini (Heteroptera: Corei- 
dae), Ph.D. Dissertation, Iowa State University, 
Ames, Iowa. 

Stal, C. 1867. Bidrag till hemipterernas systematik. 
Ofversigt af Kongliga Svenska Vetenskaps-Aka- 
demiens Forhandlingar 24 (7): 491-560. 

. 1870. Enumaeratio hemipterorum. Pt. 1. 

Svenska Vetenskaps-Akademiens Handlingar 
9(1): 1-232. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 99-105 

OBSERVATIONS ON TWELVE FAMILIES OF HOMOPTERA IN MACAU, 
SOUTHEASTERN CHINA, FROM 1989 TO THE PRESENT 

Emmett R. Easton and Wing-Wah Pun 

(ERE) University of Macau, P.O. Box 3001, Macau; (WWP) Departamento de Services 
Agrarios de Camara Municipal das Ilhas, Coloane, Macau 



Abstract. — The insect species found in present day Macao are largely those feeding 
upon small numbers of remnant agricultural plantings or on those plants representing the 
subtropical evergreen broad leaf forest or remnants of the tropical monsoon rainforest as 
agricultural crops are only grown on a small scale. Records of 37 species of Homoptera 
of the Aleyrodidae, Aphididae, Cicadellidae, Cicadidae, Coccidae, Delphacidae, Diaspi- 
didae, Flatidae, Fulgoridae, Margarodidae, Pseudococcidae and Psyllidae are listed for the 
Portuguese territory of Macao of which 27 are new records. 

Key Words: Macao, Homoptera, faunal list 



In the early part of the present century a 
number of homopteran species were listed 
as occuring in Macao and Hong Kong as 
well as part of the Guangdong Province of 
mainland China all of which was known at 
that time as South China (Kirkaldy 1909). 
Rice was considered the main agricultural 
crop and farming was the chief livelihood 
of most of the people in south east China. 
The land area that John C.W. Kershaw (Kir- 
kaldy 1909) collected insects on was con- 
sidered an island at that time but it is now 
connected to the mainland of China and to- 
day is considered the peninsula. The ma- 
jority of the collections made in this study, 
however were taken from the islands of Tai- 
pa and Coloane which are connected to the 
peninsula either by two bridges (Taipa) or 
a causeway (Coloane) which did not exist 
in the early part of the century. Many of the 
insect species noted to occur in Macao at 
the beginning of the century may now be 
non-existant. We were unable to find evi- 
dence, for example, of the planthoppers de- 
scribed by Frederick Muir (1913) because 
of modernization and construction of hotels 



and apartment complexes that have re- 
placed agricultural farmlands. Easton and 
Pun (1997) discussed the species of true 
bugs in the region and here we discuss the 
Homoptera. 

Materials and Methods 

Insects were sampled routinely from the 
walls of window-lit buildings (Taipa island) 
that are often illuminated at night. Other in- 
sect groups less sensitive to light, such as 
the cicadas other than those in the genera 
Cryptotympana, and Huechys, as well as 
the Flatidae, Fulgoridae and the scale in- 
sects, were sampled with an insect net from 
trees, shrubs and grasses in forested areas 
on the islands of both Coloane and Taipa as 
well as from the peninsular area connected 
to the Guangdong region of China. The 
names of the plant hosts follow the Macau 
Catalogue of plants and their addenda 
(1991). Voucher specimens of the Homop- 
tera named here are housed in the Ento- 
mology Museum of the Agrarian Services 
on Coloane (Seac Pai van Park) under the 
curatorship of the second author. 



100 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



List of Species 

HOMOPTERA 

Aleyrodidae 

Aleurocanthiis spiniferus (Quaintance), 
orange spiny whitefly. No date, collector 
unknown, NEW RECORD, Coloane Island, 
Macao, ex. Ficus rumphi, Rosa chinensis. 
Mound and Halsey (1978) list it from India, 
Sri Lanka, Taiwan, Thailand, Malaysia, 
Philippines, Sumatra, Japan, and Hong 
Kong, while Peng and Liu (1992) report it 
from the Fujian, Guangxi, Henan, Hunan, 
Jiangsi, and Zhejiang provinces of China. 

Trialeurodes vaporariorum (Westwood), 
greenhouse whitefly. No date, collector un- 
known, NEW RECORD. Coloane Island, 
Macao, ex. Brassica oberocea. Citrus med- 
ico and Hybiscus rosa-sinensis. Mound and 
Halsey (1978) list it from Malaya, India, Sri 
Lanka, Hawaii, New Guinea, New Zealand 
and most of the provinces of mainland Chi- 
na. 

Aphididae 

Aphis gossypii Glover, cotton or melon 
aphid. 5 Dec 1992, PWW leg. NEW RE- 
CORD, Coloane Island, ex. Althaea rosea. 
Capsicum friitesceris. Chrysanthemum mor- 
ifolium, CitruUus lanatus, Colocassia es- 
culenta, Cucurbita moschata, Dianthus car- 
yophylhis. Hibiscus rosa-sinensis, Lilium 
japonicum, Litchi chinensis, Phaseolus ra- 
diatus, Psidium guajava, Punica granatum 
and egg plant, Solanum melongena. It has 
been reported in Hong Kong (Lee and Win- 
ney 1981) and is widely distributed in Chi- 
na (Peng and Liu 1992). 

Aphis nerii Boyer de Fonscolombe, ner- 
ium or oleander aphid. 5 Dec 1992, PWW 
leg, NEW RECORD, Coloane Island, Ma- 
cau, ex. Asclepius curassavica, Nerium in- 
dicum. Raychaudhuri (1980) lists it 
throughout India and Japan, Java, Korea, 
and Taiwan. In Hong Kong, Lee and Win- 
ney (1981) report it common during winter 
months when it feeds on the shoots and 
leaves of oleander. Peng and Liu (1992) list 



it from Guangdong, Guangxi, Hunan, and 
Jiangsu provinces of China. 

Formosaphis micheliae Takahashi. 21 
March 1995, PWW leg. NEW RECORD, 
Coloane Island, Macao, ex. white jade or- 
chid tree, Michel ia alba and M. champaca. 
Blackman and Eastrop (1994) record it 
from Japan and Taiwan and Lee and Win- 
ney (1981) found it in Hong Kong. 

Myzus persicae (Sulzer), green peach 
aphid. No date or collector, NEW RE- 
CORD, Coloane Island, Macao, ex. Bras- 
sica chinensis, B. oberacea and Prunus per- 
sica. This is considered a cosmopolitan 
temperate species and it is more common 
during winter months in Hong Kong with 
populations declining after April (Hill et al. 
1982). It is widely distributed in China. 

Neophyllaphis podocarpi Takahashi, 
buddhist pine aphid. 9 Feb 1995, PWW leg. 
NEW RECORD, Coloane Island, Macao, 
ex. Podocarpus macrophyllus. Distribution 
includes Malaysia, Taiwan, Hong Kong, Ja- 
pan and the Guanxi, Hunan, Jilin, and Zhe- 
jiang provinces of China (Peng and Liu 
1992). 

Shivaphis celti B. Das. 17 March 1994, 
PWW leg. NEW RECORD, Coloane Is- 
land, Macao, ex. Celtis sinensis. Distribu- 
tion includes Korea, Japan, India, Taiwan, 
Hong Kong and the Fujian, Guangdong, 
Guangxi, Guizhou, Hebei, Hunan, Jiangsu, 
Liaoning, Shandong, Sichuan, and Yunnan 
provinces of China (Peng and Liu 1992). 

Tinocallis kahawaluokalani (Kirkaldy), 
crepe myrtle aphid. 27 Oct 1993, PWW leg. 
NEW RECORD, Coloane Island, Macao, 
ex. Lagerstroemia indica. This species is 
also found in India, Taiwan, Japan, and Ha- 
waii (Raychaudhuri 1980). 

Cicadellidae 

Lodiana brevis (Walker), yellow-banded 
leafhopper. 9 Dec 1993. Ng Wai Man leg. 
NEW RECORD, Coloane Island, Macao, 
ex. Cinnamomum camphora, Ficus micro- 
carpa and Euphoria longan. In Hong Kong 
Lee and Winney (1981) reported it from 
Citrus sp. Distribution includes the Guang- 



VOLUME 101. NUMBER 1 



101 



dong, Guizhou, and Yunnan provinces of 
China as well as India, Taiwan, Malaysia, 
Thailand, and Japan (Datta 1988). 

Nephotettix virescens (Distant), green 
rice leafhopper. No date or collector, NEW 
RECORD, Coloane Island, Macao, ex. Cit- 
rus sp., Bambusa sp., and Sacchanim offi- 
cinarum. There are records of it from rice 
in the Tai Lung farm insect collection in 
Hong Kong and Hill (1975) lists it as a pest 
of rice in India and China. 

Petalocephala chlorocephala Walker. 4 
Sept 1993, Ng Wai Man leg, NEW RE- 
CORD, Coloane Island, Macao, no host 
data. 

Tettigoniella spectra (Distant), rice white 
leafhopper. 15 Sept 1992, Ng Wai Man leg, 
NEW RECORD, Coloane Island, Macao, 
ex. Morns alba, Oryza sativa, Saccharum 
officinahum. Distribution includes Taiwan, 
India, and Japan as well as the Guangdong 
and Hunan provinces of China (Peng and 
Liu 1992). 

Another species of leafhopper, Dryado- 
morpha pallida Kirkaldy has not been col- 
lected in our study but Webb (1981) re- 
ported it from Macao and Hong Kong as 
well as the neighboring areas of India, 
Bangladesh, Taiwan, Japan, Laos, and the 
Philippines. 

Cicadidae 

Cryptotympana atrata (E), large brown 
cicada. 20 June 1994. Cheong Chi Kong 
leg, Coloane island, Macao, ex. Melia aze- 
darach. Moms alba, Tectonis grandis. 
Emergence of adults occurs from late May 
through early July in general but emergence 
on Taipa Island in 1998 was earlier (May 
15). Distribution includes Hong Kong, the 
provinces of Guangdong, Hebei, and Zhe- 
jiang of China (Wu 1935) as well as Ma- 
laysia, Japan, and Taiwan. 

Cryptotympana mandarina Distant. May 
1997, 15 May 1998, ERE leg, Taipa Island, 
Macao. Kershaw (1903) is believed to be 
the first to record it from Macao while Kir- 
kaldy (1909) reported it from Hong Kong. 
Nymphal skins have been observed on the 



trunks of various trees such as Acacia con- 
fusa and Casuarina equisetifolia in the mu- 
nicipal cemetary on Taipa island (Easton 
1992) as well as the foxglove tree, Paulow- 
nia fortunei on Guia Hill of Macao penin- 
sula in May, 1997 and on Hibiscus tiliaceus 
near the Monte Forte, Fortress, May 10, 
1998. Emergence holes near the base of the 
trees indicated where the nymphs left the 
soil. Watery fluid has been observed emit- 
ting from the anal region of resting adults 
on Taipa Island suggesting that they had re- 
cently emerged from the ground. As feed- 
ing in adults has not been documented, the 
watery fluids may have accumulated while 
the nymphs were underground and release 
may be necessary before sound production 
can take place. Distribution also includes 
the Guangdong and Hainan provinces of 
China and Taiwan. 

Chremistica ochracea (Walker). 23 May 
1995, 10 May 1998, ERE leg, Taipa Island, 
Macao. Hayashi (1977) recorded this spe- 
cies from Macao, and Hill et al. (1982) il- 
lustrated it as the green clearwing cicada 
under the genus Dundubia in Hong Kong. 
Distribution also includes Taiwan and the 
Guangdong region of China. 

Gaeana maculata Walker, yellow-spotted 
black cicada. 14 April 1994, Ng Wai Man 
leg, Coloane Island; 7 May 1998, ERE leg, 
Guia Hill, Macao peninsula. In the Seac Pai 
Van agricultural park on Coloane Island, 
nymphs were observed emerging as early 
as March 25 (Easton 1992). It has not been 
observed on Taipa island suggesting certain 
food plants may be necessary for nymphal 
development. In Hong Kong, Cheung and 
Marshall (1973) reported it from Schiruis 
terebinthiofolius, Christmas berry tree, and 
Paulownia tonnetosa and in Macao the se- 
nior author has observed it among the fo- 
liage of Pawlownia fortunei on the Guia 
Hill. Wu (1935) recorded it from Myanmar 
(Burma), Assam area of India, Vietnam, 
and Guangxi Province of China. 

Huechys sanguineus (DeGeer), red-nosed 
cicada. 31 August 1992, Ng Wai Man leg, 
Coloane Island, Macao. Kirkaldy (1909) 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



first reported this species from Macao and 
Hong Kong. From 1989-1997 emergence 
from the ground has always been later in 
the year than with the former species and 
from September through December (Easton 
1991). In 1998 however an emergence of 
15-20 individuals was observed in an ar- 
boretum on Coloane island on May 10. 
Specific food plants have not been associ- 
ated with nymphal feeding. 

Mogannia hebes (Walker), grass cicada. 
9 June 1994, ERE leg, NEW RECORD, 
Taipa Island on shrub on university campus; 
June 1996, May 1998, ERE leg, Seac Pai 
Van Agricultural Park, Coloane Island. Kir- 
kaldy (1909) was probably the first to re- 
cord it from Hong Kong, and Hayashi 
(1976) described its feeding on the stems of 
Miscanthus grass where females oviposit 
into stems and leaf midribs from April to 
June. Distribution includes Korea, India, 
Myanmar, and the Chejiang, Fujian, Guang- 
dong, Guangxi, Jiangsi, Hunan, Sichuan, 
and Yunnan provinces of China. 

Mogannia nasalis (White). 25 May 1966, 
ERE leg, Taipa Island, Macao on tree fo- 
liage along hiking trail in thickly forested 
hillside. This species was first recorded in 
Macao and Hong Kong by Kirkaldy (1909). 
It is also found in the Assam area of India. 

Platypleura hilpa Walker, spotted brown 
cicada. 2 August 1992, PWW leg, Coloane 
Island, Macao. Nymphs were observed 
emerging at night near the trunks of Casu- 
arina equisetifolia trees near the municipal 
cemetary on Taipa Island (Easton 1992). 
Kirkaldy (1909) reported it from Hong 
Kong and Macao, and it is also found in the 
Guangdong and Hunan provinces of China 
(Peng and Liu 1992). 

Coccidae 

Ceroplastes cerifenis (Fabricius), indian 
wax scale. 18 Dec 1992. PWW leg, NEW 
RECORD, Coloane Island, Macao, ex. Cin- 
namomum camphora, Melastoma sangiii- 
neum, Michelia figo. Moms alba. Tang 
(1991) listed it from Australia and Hawaii 
and it is also found in Japan, India, Sri Lan- 



ka and provinces of China south of the 
Yangtze River. 

Saissetia cojfeae (Walker), helmet or 
hemispherical scale. 19 Jan 1988, PWW 
leg, NEW RECORD, Coloane Island, Ma- 
cao, ex. Fukien tea, Carmona microphylla 
and sago palm, Cycas revoluta. Hill (1975) 
listed citrus, guava and mango as alternate 
hosts but mainly it is found on coffee plants 
and widespread in the tropics including 
southeastern Asia. 

Delphacidae 

Nilaparx'ata lugens (Stal), brown plant- 
hopper of rice. 22 Oct 1997, ERE leg, 
NEW RECORD, Taipa Island on window 
ledges and outside on floors of university 
campus buildings illuminated by lights. Ac- 
cording to Hill (1975) and Wada et al. 
(1987), N. lugens invades Japan with the 
monsoon winds every year from China, so 
the large numbers (1,000 + ) observed in 
Macao over a brief period of 1-2 days sug- 
gests that the insects migrated from the 
neighboring Guangdong Province since rice 
is not grown commercially locally in Ma- 
cao as a crop. In Malaysia and Indonesia, 
it is a major pest of rice and produces a 
browning effect on the plants known as 
"hopperburn." Lee and Winney (1981) also 
report it from Gladiolus gandavensis in 
Hong Kong. Yang (1989) gives its distri- 
bution as Australia, New Guinea, India, Ko- 
rea, Taiwan, Philippines, Vietnam, and the 
Pacific islands of Fiji, Guam, Yap, and Pa- 
lau, while Kuoh et al. (1983) list it from 
Anhui, Fujian, Gansu, Guangdong, Gu- 
angxi, Guizhou, Hebei, Henan, Honan, Hu- 
nan, Jiangsu, Jiangxi, Jilin, Liaoning, Shan- 
dong, Shanxi, Yunnan, and Zhejiang Prov- 
inces in China. 

Other species of Delphacidae reported 
earlier by Muir (1913), such as Belocera 
sinensis Muir, Phyllodinus macaoensis 
Muir, and Tropidocephala saccharivorella 
Matsumura, were not collected in this study 
and may no longer be found locally due to 



VOLUME 101. NUMBER 1 



103 



the urbanization which has replaced agri- 
cultural crops. 

Diaspididae 

Aiilacaspis rosarum Borchsinius, Asiatic 
rose scale. No date or collector, NEW RE- 
CORD. Coloane Island, Macao, ex. Rosa 
chinensis. Distribution includes the Fujian, 
Guangdong, Guangxi, Hunan, Jiangsu, Ji- 
angxi, Shandong, Sichuan, Yunnan, and 
Zhejiang provinces of China (Peng and Liu 
1992). 

Aulacaspis yabunikkei Kuwana. 21 April 
1988, PWW leg, NEW RECORD, Coloane 
Island, Macao, ex. Cinnamomum campho- 
ra. In Hong Kong, Lee and Winney (1981) 
also record it from pond spice, Litsea glu- 
tinosa. Distribution includes Japan and the 
Guangdong, Guizhou, Hunan, Sichuan, 
Yunnan, and Zhejiang Provinces of China 
(Peng and Liu 1992). 

Hemiberlesia pitysophila Takagi. 7 June 
1988, PWW leg, NEW RECORD, Coloane 
Island, Macao, ex. Pinus mansoniana. 

Lepidosaphes laterochitinosa Green. 20 
May 1997, PWW leg, NEW RECORD, Co- 
loane Island, Macao, ex. Osmanthiis fra- 
grans, Kwai-Fah. In Hong Kong there are 
records of it from this host as well as Schef- 
flera octophylla. Ivy tree, in the Tai Lung 
Farm insect collection. 

Parlatoha pergandii Comstock, chaff 
scale. 23 Feb 1993, Yau H.C. leg, NEW 
RECORD, Coloane Island, Macao, ex. Jas- 
mmiiim sambac and Citrus sp. Distribution 
in the world is widespread, but in Asia it 
has been recorded from Australia, New 
Zealand, Japan, India, Philippines, Taiwan, 
and from the Anhui, Fujian, Guangdong, 
Guangxi, Hainan, Hebei, Henan, Hubei, 
Hunan, Jiangsi, Jiangsu, Liaoning, Qinghai, 
Shanxi, Shaanxi, Shanghai, Sichuan, Yun- 
nan, and Zhejiang Provinces of China 
(Peng and Liu 1992). 

Pseudaulacaspis cockerelli (Cooley), 
oyster or oleander scale. 7 May 1988, 
PWW leg, NEW RECORD, Coloane Is- 
land, Macao, ex. Michelia alba, M.figo and 
coconut palm, Cocos nucifera. In Hong 



Kong, there are also records from bamboo 
palm, Chrysalidocarpiis lutescens (Lee and 
Winney 1981). Its distribution includes 
Thailand, Taiwan, and the Guangdong, Hu- 
bei, Hunan, Jiangsu, Jiangxi, Shandong, 
Sichuan, Yunnan, and Zhejiang provinces 
of China (Peng and Liu 1992). 

Flatidae 

Lawana imitata (Melichar), white moth 
bug. 18 July 1991, PWW leg, Coloane Is- 
land, Macao, ex. Bauhinia sp.. Euphoria 
longan. Hibiscus tiliaceus, Jasminium mes- 
nyi, Litsea monopetala, Murraya panicula- 
ta, and Pittosporum tobira. This insect is 
very common in lychee fruit tree orchards 
near Shenzhen in the Guangdong region of 
China. Easton (1992) reported it from the 
Seac Pai Van Agricultural Park in Macau. 
Its distribution also includes the Hainan, 
Hunan, Guizhou, Guangxi, and Yunnan 
provinces of China, Japan (Peng and Liu 
1992) as well as Hong Kong (Lee and Win- 
ney 1981). 

Seliza lignaria (Walker). 9 May 1997, 
ERE leg, NEW RECORD, Taipa Island, 
Macao, ex. Miscanthus grass along hiking 
trail in recently burned vegetation. Fennah 
(1956) listed it from Hong Kong and the 
Guangdong province of China, and it is also 
believed to occur in the Anhui, Fujian, 
Guizhou, Hunan, Sichuan, Yunnan, and 
Zhejiang provinces of China and India 
(Peng and Liu 1992). 

Medler (1992) reported Salurnis margi- 
nella (Guerin-Meneville) and Geisha dis- 
tinctissima (Walker) as occuring in Macao 
in the early part of this century, but we have 
been unable to document their presence in 
this study. 

Fulgoridae 

Fulgora candelaria (L.), lantern fly or 
lantern bug. 2 May 1994, Cheong Pak Fai 
leg, Coloane Island; 10 Dec 1990, ERE leg, 
Guia Hill near lighthouse, Macao Peninsu- 
la, ex. Litchi chinensis. Euphoria longan, 
Morus alba, Sapium sebiferum. Kershaw 



104 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



and Kirkaldy (1910) were probably the first 
to describe, illustrate, and discuss the life 
history stages in the area known as South 
China which includes Hong Kong and Ma- 
cao. Even though four host plants are given 
above, Kershaw and Kirkaldy (1910) felt 
that E. longan and mango trees were the 
most important for the reproduction of this 
species. Its distribution also includes India 
(Assam), Cambodia and the Hainan, Hu- 
nan, Guangdong, Guangxi, and Sichuan 
provinces of China. 

Zanna chinensis (Distant). 31 Aug 1990, 
ERE leg, Taipa Island, Macao, ex. Casua- 
hna equisetifolia, horsetail tree. There are 
also records of this species in Hong Kong 
from the Tai Lung Farm insect collection. 
Wu (1935) recorded it from the Naga Hills 
area between India and Myanmar (Burma). 

Margarodidae 

leery a purchasi Maskell, cottony cushion 
scale. 10 Feb 1994, PWW leg, Coloane Is- 
land; 12 May 1997, ERE leg, Taipa Island, 
NEW RECORD, ex. Cassia surattensis, the 
sunshine tree, Acaeia eonfusa, Aealypha 
wilkesiana, Casuarina equisetifolia, Hyper- 
ieum ehinense, Chinese St. Johnswort, and 
Pentas laneeolata. In Hong Kong, Lee and 
Winney (1981) reported this species from 
Rosa spp. and Citrus litnonia, Chinese lem- 
on. It has a widespread distribution includ- 
ing Japan, Korea, Philippines, Indonesia, 
Malaysia, and Sri Lanka in southeastern 
Asia, including the Gansu, Guizhou, Hei- 
longjiang, Hunan, Ningxia, Qinghai, and 
Xinjiang provinces of China (Peng and Liu 
1992). 

Pseudococcidae 

Maeonellieoeeus hirsutus (Green), Asian 
hibiscus mealybug. 16 June 1988, PWW 
leg, NEW RECORD, Coloane Island, Ma- 
cao, ex. Hibiseus rosa-sinensis. In Hong 
Kong, it has been reported from Cuban 
bast, H. tiliaceus. Williams (1996) listed it 
from Bangladesh, Brunnei, Myanmar (Bur- 
ma), Cambodia, India, Indonesia, Laos, 
Malaysia, Nepal, Pakistan, Philippines, Sri 



Lanka, Thailand, and China in southeastern 

Asia. 

Psyllidae 

Macrohomotoma striata Crawford, fig 
shoot psyllid. 29 Dec 1993, PWW leg, 
NEW RECORD, Coloane Island, Macao, 
ex. Fieus microcarpa and Fieiis retusa. In 
Hong Kong, Hill and Cheung (1978) de- 
scribed small waxy colonies inhabiting api- 
cal and lateral shoots of twigs and branches. 
Hollis and Broomfield (1989) reported it 
from India and Hodkinson, (1986) listed it 
from Ryukyu Is. 

In the family Tropiduchidae Lee and 
Winney (1981) reported Kallitaxila ma- 
eaoana (Muir) in Hong Kong, but we have 
not collected it in this study even though 
the original type locality was in Macao 
(Muir 1913). 

Acknowledgments 

The authors thank Clive S.K. Lau and 
Stephen Lai Yue-hong, agricultural officers. 
Hong Kong Government, for allowing one 
of us (ERE) access to their insect collection 
on the Tai Lung Farm near Sheung Shui, 
New Territories. We are also grateful for 
species determinations of the Flatidae by 
John T Medler of the B.P. Bishop Museum, 
Honolulu, Hawaii, and to J. P. Duffels, Zool- 
ogisch Museum, Instituut voor Systematiek 
en Populatiebiologie, Amsterdam, The 
Netherlands, for determination of the cicad- 
idae. The delphacid planthopper was iden- 
tified by Yang Jeng-Tze of the Department 
of Entomology, National Chung Hsing Uni- 
versity, Taichung, Taiwan. 

Literature Cited 

Blackman, R. L. and V. F. Eastrop. 1994. Aphid's on 
the World's Trees. An Identification and Infor- 
mation Guide, CAB International, Natural History 
Museum, London. 986 pp. 16 pis. 

Cheung, W. W. K. and A. T Marshall. 1973. Water 
and ion regulation in cicadas in relation to xylem 
feeding. Journal of Insect Physiology 19: 1801- 
1816. 

Datta. B. 1988. On Oriental cicadellidae (Homoptera: 
Insecta). Records of the Zoological Society of In- 



VOLUME 101, NUMBER 1 



105 



dia. Miscellaneous Publications, Occasional Paper 
No. 90. 256 pp. 

Easton, E. R. 1991. Annotated list of the insects of 
Macau observed during 1989. Entomological 
News 102(2): 105-111. 

Easton, E. R. 1992. 1990 additions to the annotated 
list of the insects of Macau. Entomological News 
103(1): 30-36. 

Easton, E. R. and W. W. Pun. 1997. Observations on 
some Hemiptera/Heteroptera of Macau, southeast 
Asia. Proceedings of the Entomological Society of 
Washington 99(3): 574-582. 

Fennah, R. G. 1956. Fulgoroidea from southern China. 
Proceedings of the California Academy of Scien- 
ces 28(13): 441-527. 

Hayashi, M. 1976. On the species of the genus Mo- 
gannia Amyot et Serville 1853 (Homoptera, Ci- 
cadidae) of the Ryukyus and Taiwan. Kontyij 
44(1): 27-42. 

Hayashi, M. 1977. New records of cicadas (Homop- 
tera, Cicadidae) from Hong Kong. Kontyu 45(1): 
91. 

Hill. D. S. 1975. Agricultural Insect Pests of the Trop- 
ics and Their Control. Cambridge University 
Press, Cambridge. U.K., 746 pp. 

Hill, D. S. and W. W. K. Cheung. 1978. Hong Kong 
Insects. Urban Council, Hong Kong, 128 pp. 

Hill, D. S., P Hore, and I. W. B. Thornton. 1982. In- 
sects of Hong Kong. Hong Kong University Press, 
Hong Kong, 503 pp. 

Hodkinson, I. D. 1986. The Psyllids (Homoptera: Psyl- 
loidea) of the Oriental zoogeographical region: An 
annotated checklist. Journal of Natural History 
20(2): 299-357. 

Hollis. D. and P S. Broomfield. 1989. F/c//i-feeding 
psyllids (Homoptera) with special reference to the 
Homotomidae. Bulletin of the British Museum of 
Natural History (Entomology) 58(2): 131-183. 

Kershaw, J. C. W. 1903. A naturalist's notes from Chi- 
na. Field Naturalist Quarterly 2: 233-235. 

Kershaw, J. C. W. and G. W. Kirkaldy. 1910. A mem- 
oir on the anatomy and life-history of the homop- 
terous insect Pyrops ccmdelaria (or candle-fly) 
Zoologische Jahrbucher 29(2): 105-124. 

Kirkaldy, G. W. 1909. A list of the Hemiptera of ori- 
ental China. Part I. Annales de la Societe Ento- 
mologique de Beligique 53: 177-183. 

Kuoh, C.-L., Ding, J.-H.. Tian Li-X., and C. L. Hwang. 
1983. Homoptera: Delphacidae. Economic Insect 
fauna of China. Fascicle 27. Science Press, Bei- 
jing 165 pp. 13 pis. (In Chinese.) 

Lee. L. H. Y. and R. Winney. 1981. Checklist of ag- 



ricultural insects of Hong Kong 1981. Agriculture 
and Fisheries Department Bulletin Number 2, 
Hong Kong, 164 pp. 

Macau. Catalogue of the Plants of Macau. 1991. Ca- 
mara Municipal das Ilhas. Instituto de Investiga- 
gao Cienti'fica Tropical, Lisboa. 177 pp. 

Medler, J. T 1992. Revision of the tribe Phyllphantini 
in the Oriental Region with descriptions of new 
genera and new species (Homoptera:Flatidae) Ori- 
ental Insects 26: 1-38. 

Mound, L. A. and S. H. Halsey. 1978. Whitefly of the 
World, A systematic catalogue of the Aleyrodidae 
(Homoptera) with host plant and natural enemy 
data. British Museum (Natural History) and John 
Wiley and Sons, Chichester. 340 pp. 

Muir, F. 1913. On some new fulgoroidea. Proceedings 
of the Hawaiian Entomological Society 2(5): 237- 
269. 

Peng, J.-W. and Liu, Y.-Q. 1992. Editors in Chief. Ico- 
nography of Forest Insects in Hunan, China. In- 
stitute of Forestry Sciences, 1,473 pp. (In Chi- 
nese.) 

Raychaudhuri, D. N. 1980. Aphids of North-east India 
and Bhutan. The Zoological Society, Calcutta. In- 
dia. 520 pp. 

Tang, F-D. 1991. The Coccidae of China. Shanxi Unit- 
ed Universites Press, Taiyuan, China, 377 pp, 84 
pis. (In Chinese.) 

Wada, T, H. Seino, Y. Ogawa, and T Nakasuga. 1987. 
Evidence of autumn overseas migration in the rice 
planthoppers, Nilapan'ata liigens and Sogatella 
furcifera: analysis of light trap catches and asso- 
ciated weather patterns. Ecological Entomology 
12: 321-330. 

Webb, M. D. 1981. The Asian, Australasian and Pa- 
cific Paraboloponinae (Homoptera: Cicadellidae), 
a taxonomic revision with key to all the known 
genera of the subfamily. Bulletin of the British 
Museum of Natural History (Entomology) 43(2): 
1-76. 

Williams, D. J. 1996. A brief account of the hibiscus 
mealybug, Maconellicoccus hirsutus (Hemiptera: 
Pseudococcidae), a pest of agriculture and horti- 
culture, with descriptions of two related species 
from southern Asia. Bulletin of Entomological 
Research 86: 617-628. 

Wu, C. F 1935. Catalogus Insectorum Sinensium (Cat- 
alogue of Chinese Insects) II. The Fan Memorial 
Institute of Biology, Beijing, China, 634 pp. 

Yang, C. T. 1989. Delphacidae of Taiwan (II) (Ho- 
moptera: Fulgoroidea) National Science Council 
Special Publication No. 6, Republic of China, 
334 pp. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999. pp. 106-122 

BIOLOGY OF ANDRENA (SCRAPTEROPSIS) FENNINGERI VIERECK 
(HYMENOPTERA: ANDRENIDAE), HARBINGER OF SPRING 

Suzanne W. T. Batra 

Bee Research Laboratory, PSI, Agricultural Research Service, U.S. Department of Ag- 
riculture, Bldg. 476, BARC-East. Beltsville, MD 20705 



Abstract. — Andrena fenningeh Viereck is the first native bee species to fly each spring 
at Beltsville, MD; males appear as early as February 9, sometimes before their floral hosts 
begin to bloom. A permanent aggregation of nests in red clay was studied for 10 years. 
These univoltine, solitary bees break diapause and move at 4°C from their natal cells 
toward the soil surface in midwinter, ready to emerge and mate as soon as the topmost 
soil warms. They thermoregulate by aggregating their nests in the warmest available 
microclimate, and by basking on cold, sunny days to achieve the minimum 1 TC required 
for flight. The most important host is Acer rubrum; Primus, Pyrus, and Salix are also 
visited. Male swarming behavior, phenology, nest structure, and associates (including 5 
Nomada spp., Myopa sp., and the behavior of 3 unusual species of Eiistalomyia [Antho- 
myiidae]) are discussed. This bee may be manageable as an orchard pollinator, if suitable 
microhabitat and supplemental hosts are provided. 

Key Words: bees, nests, thermoregulation, phenology, fruit pollination, Eiistalomyia, 
conopids, Acer 



The Holarctic bee genus Andrena in- 
cludes about 700 Eurasian and 500 North 
American species; both species and individ- 
uals are dominant components of the bee 
fauna during springtime; they are important 
pollinators of crops and wild plants (see ref- 
erences in Batra 1990). The behavior and 
ecology of few species have been studied. 
Most species of Andrena are solitary, uni- 
voltine, polylectic bees; many of them share 
hosts, geographic ranges, and times of adult 
activity; the reasons why there are so many 
species of Andrena remain unknown. 

This report concerns the nesting behavior 
of an Eastern, cold-adapted, polylectic, sol- 
itary species, which is the first native bee 
species to fly each spring at the Beltsville 
Agricultural Research Center (Prince 
George's Co., Maryland). Adult activity of- 
ten begins while snow and ice remain on 



the frozen ground in shady places, and no 
plants are yet in bloom. Andrena fenningeri 
Viereck (det. W. E. LaBerge) is in the North 
American subgenus Scrapteropsis, which 
includes 18 vernal species (LaBerge 1971). 
The nesting behavior of only one of them, 
A. (Scrapteropsis) alleghaniensis Viereck, 
has previously been studied (Batra 1990). 
This solitary bee resembles A. fenningeri in 
its preference for Acer as a nectar and pol- 
len source and in the location of its nest 
aggregation so as to maximize insolation. 

I investigated the behavior of A. fennin- 
geri at intervals over a period of 10 years 
(1987 to 1997), at an aggregation of nests 
in a sunny spot at the north edge of a large 
field (the "Rose Garden," off Entomology 
Rd.). Aggregations of nests of Andrena 
may persist for decades (Chambers 1968, 
Schonitzer and Klinksik 1990, 1992, Rid- 



VOLUME 101. NUMBER 1 



107 



dick 1992). If the factors that permit or en- 
courage permanent aggregations could be 
determined, this basic information may 
prove useful in conserving existing natural 
aggregations, and also for actively manag- 
ing Andrena bees to pollinate fruit crops. 
For these reasons, the unusually early adult 
activity of A. fenningeri was investigated. 

Materials and Methods 

I made observations early in each season 
and during fair weather, when bees were 
flying. A total of 214 hours were spent, 
most of them during the unusually warm 
and early spring of 1990 (76 hours), and the 
cool, late spring of 1992 (60 hours). En- 
trances to nests with tumuli were marked 
with small, numbered aluminum tags, in- 
serted 2 cm north of each nest. Meteorolog- 
ical data were recorded during visits. Soil 
temperatures at sites 1-6 were taken by in- 
serting calibrated bimetallic dial probe ther- 
mometers to the appropriate depths. Nests 
were excavated by shovel and trowel after 
pouring plaster into them, which rendered 
tunnels easily visible. Adult bees and clep- 
toparasites were netted, then pinned, or pre- 
served in FAA for dissection under a mi- 
croscope. Pollen was stained with lacto- 
phenol-cotton blue and examined micro- 
scopically. I indicate means after ranges (in 
parentheses). Voucher specimens will be 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, D. C. 

Nests and their Contents 

The nests of A. fenningeri in level ground 
(Fig. 1) were vertical tunnels that penetrat- 
ed the uppermost humus-and-root-filled 
zone of the clay soil, to a zone of dense, 
poorly drained, red marine clay, where the 
cells were made. Nests ended at the top of 
a zone of gritty "hardpan." From late fall 
through early spring, the red clay was very 
moist, often muddy; after the tree canopy 
had leafed out in May (after adult bees 
ceased activity), the upper part of the clay 
dried to an adobe-like, hard consistency 



(soil penetrometer reading 3.5-4.5 kg/cm), 
providing protection to the growing brood 
and dormant adults in the cells. Evaporation 
and transpiration by plants during the hot 
and relatively dry summer months dried the 
clay. Rainy, cool weather, and the cessation 
of most transpiration after November per- 
mitted the soil to moisten and soften. Adult 
bees emerged from their cells in the soft- 
ened soil, and crawled toward the surface 
of the soil during mid-winter. They waited 
just below the soil surface by late winter, 
ready to fly during the first warm days of 
spring. 

A total of 22 entire nests was examined 
(5 in 1987; 3 in 1989; 14 in 1991). Because 
the lateral tunnels that led to cells were 
backfilled by the solitary mother bees with 
soil and obliterated after oviposition, many 
cells that were found could not be traced to 
their nests' tunnels (Fig. 1) The nests' tun- 
nels reached depths of 16-25 (20.8) cm, 
and were 4.5-5.5 mm in diameter. Nest en- 
trances were irregular, 3.5-5.5 mm in di- 
ameter, and when new, surrounded by a 
usually circular tumulus of loose, dry, soil 
particles 2.0-5.0 (3.3) cm in diameter and 
0.5-1.0 cm high. The tumulus may not be 
rebuilt when it had disappeared due to rain. 
Nest entrances were often closed by loose 
soil particles when the bees were not for- 
aging. The entrances to about 20% of the 
nests, which had been initiated beneath fall- 
en leaves or in dense turf grasses, were dif- 
ficult to find. 

Nests had up to 5 brood cells (1.5) each. 
They were at the ends of lateral tunnels, 
which were 1.0-6.0 (2.5) cm long and 3.5- 
5.0 mm in diameter. Cells were made at 
depths of 13.0-26.0 (21.4) cm. The 39 cells 
were nearly horizontal, of the usual ovoid 
shape of Andrena cells, and coated inter- 
nally with thin, shining, transparent water- 
proof linings, secreted by Dufour's glands. 
Cells were 10.0-12.5 (1 1.0) mm long, 5.0- 
6.0 (5.5) mm in maximum diameter, taper- 
ing to the cell's entrance, which was 2.5- 
4.0 (3.3) mm in diameter. After oviposition, 
this entrance was sealed with compressed 



108 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Two nests of A. fenningeri in red marine clay soil(stippied area). Nests extended to a zone of denser 
soil (hardpan, hatched area), at a depth of 23 cm. The lateral tunnels that led to completed cells were filled with 
soil by the mother bees. Nest A, closed at the surface, was without an adult bee. It had 3 cells with eggs on 
pollen balls (examined May 4, 1989). Nest B was open, but without a tumulus. Two sealed cells contained eggs 
on pollen balls; one of these cells (C) had droplets of moisture on its thin, smooth waterproof lining. Another, 
open cell (D) had a dead female bee with a large, live conopid maggot filling her abdomen. She died in the 
position taken by bees when they oviposit (nest examined May 7, 1987). 



VOLUME 101, NUMBER 1 



109 



soil pellets, laid down in two concentric 
rings around a central depression. The Du- 
four's-gland secretion in many genera of 
bees varies in composition. It polymerizes, 
forming a thin, smooth solid that water- 
proofs subterranean cells; it can also func- 
tion as a pheromone, and as food for larvae. 
In Andrena, it is composed mainly of iso- 
prenoid esters (see review in Ayasse et al. 
1990). 

Small, regularly-spaced, tasteless (to me) 
droplets, probably of water, were some- 
times found on the hydrophobic cell lin- 
ings, near the entrances of both empty and 
provisioned cells (Fig. IC). Perhaps the 
shape of cells influences the condensation 
pattern, keeping water from condensing on 
the hygroscopic provisions, where it would 
permit spoilage. Yeasts in nectar and am- 
bient fungi ferment and spoil the wetted 
provisions of many subterranean-nesting 
bees, causing significant mortality of their 
larvae (Batra 1970). The soil that seals en- 
trances to cells and laterals of A. fenningeri 
is without a visibly water-repellent lining. 
It is porous, permitting some circulation of 
the water-saturated subsoil air. The survival 
of this bee, living inside cells for many 
months, and the preservation of its hygro- 
scopic provisions inside humid cells, may 
depend on such a condensation-site-con- 
trolling feature of its brood cells. 

Pollen balls (provisions) of A. fenningeri 
that had been made in early spring were 
olive-green, and composed solely of pollen 
of Acer rubriim L., mixed with nectar. 
Those made after Acer bloom were various 
shades of yellow, and made of up to 3 spe- 
cies of pollen. Pollen balls were firm, 
smooth, spheroid, somewhat flat on top, 
and 3.7-4.0 mm high and 4.0-4.8 mm wide 
(Fig. IC). Occasionally, laterals and cells 
with large pollen balls, but no eggs or lar- 
vae, had been filled with earth. Eggs were 
strongly arched, white, 2.0-2.7 mm long 
and 0.5 mm thick. Larvae, ranging from 
2.5-4.0 mm long, fed on top of the provi- 
sions; they later lay on their backs beneath 
their pollen balls, and reached 8 mm long 



when all their provisions had been eaten. 
Larvae transformed to prepupae in late 
May, after defecation. No cocoons were 
made. The times of pupation and transfor- 
mation to adults were not studied. 

The time of adult emergence each spring, 
foraging, oviposition, and larval develop- 
ment varied with weather conditions each 
year. Although individual females were not 
marked, many of their nests had been. The 
appearance of new tumuli late in spring 
where there had been no nests, the mid-sea- 
son closure of nests that had been tagged in 
early spring, and the few cells per nest, in- 
dicated that some female A. fenningeri 
make more than one nest each, as do some 
other species of Andrena. At the end of the 
nesting season, during the first week of 
May, the old females became disoriented 
and exhibited displacement activity, such as 
random digging, as is seen in Andrena 
nycthemera Imhoff (Schonitzer and Klink- 
sik 1990). 

Floral Hosts 

Andrena fenningeri is polylectic (La 
Berge 1971, Hurd 1979), visiting Acer, 
Primus, Pyrus, Salix, and several other 
hosts. The first host to bloom each spring 
and the first to be visited for nectar and pol- 
len at Beltsville was Acer riibrwn. This is 
a dominant tree, occupying about 20% of 
the forest canopy near the nests. It is an 
important food resource for a wide variety 
of insects, available just as the insects 
emerge from hibernation (Batra 1985). The 
first provisions made by A. fenningeri were 
composed of the olive-green pollen from A. 
rubrum. In this respect, A. fenningeri re- 
sembles a related species, A. (Scrapterop- 
sis) alleghaniensis Viereck, which provi- 
sions its cells with maple pollen in New 
York (Batra 1990). Several species of ma- 
ple trees provide nectar and pollen for var- 
ious other Andrena bees in Europe (Cham- 
bers 1968) and North America (LaBerge 
1971; more references in Batra 1990). After 
the red maples finished blooming, A. fen- 
ningeri visited flowers of Primus, Pyrus 



110 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



and forbs, growing near the aggregation. 
Thus, the provisions that were made later 
in spring were yellow. 

The time of emergence of adult A. fen- 
ningeri in some years coincided with the 
beginning of bloom of A. rubrum and Salix 
sp. In other years, the bees emerged a few 
days before any host plants had started to 
bloom. They mated, and began nesting, 
without having eaten anything. For exam- 
ple, 2 of 3 females that were starting to ex- 
cavate nests on February 28, 1992, were in- 
seminated, but all 3 had empty crops; 14 of 
15 males in a mating swarm on that day 
had empty crops (one had eaten some ma- 
ple pollen); all of these 18 early bees had 
large fat bodies in their abdomens (meta- 
somas) that fueled their flights. By March 
2, 1992, 5 males in a mating swarm had 
empty crops and small fat bodies; 7 females 
collected while flying over the aggregation 
still had large fat bodies, but they also had 
filled their crops exclusively with A. ru- 
brum pollen, and eggs were developing in 
their ovaries. The beginning of flowering 
by A. rubrum varies by up to 2 months with 
weather conditions in early spring; it begins 
with male flowers and ends about 2-3 
weeks later, with female flowers, the trees 
being usually dioecious and dichogamous. 
Bloom began on the following dates at 
Beltsville: April 2, 1978; March 19, 1982; 
March 6, 1983 (Batra 1985); February 1, 
1989; February 12, 1990; February 21, 
1991; February 28, 1992; March 24, 1993 
and March 13, 1994. 

Thermoregulation by Aggregation 

Andrena fenningeri is unusual in its use 
of dense red clay as a nesting substrate, thus 
resembling A. macro (Riddick 1990, 1992). 
It nested in clay, even though a large area 
of exposed, sunlit sandy soil was within 
about 100 m. Most species of Andrena nest 
in more porous, well-drained soils, espe- 
cially sand (Miliczky and Osgood 1995, 
Batra 1990); hence they are called "Sand- 
bienen" in German (Gebhardt and Rohr 
1987). 



On April 21, 1987, when I discovered the 
aggregation, nests occupied an area of 5 X 
8 m. Most nests (53, with tumuli) were in 
a 3 X 3 m area with exposed soil; it was a 
buck (deer) scrape, which was renewed ev- 
ery autumn (Fig. 2, site 1). There were 18 
nests/m-, with a minimum internest distance 
of 1.0 cm. In the part of the aggregation 
that was in soil covered with short turf (site 
2), nests were fewer, spaced up to 1 m 
apart. On March 8, 1990, the aggregation 
measured 5 X 34 m; there were up to 54 
nests/m- in exposed soil (site 1), with in- 
ternest distances of 4-18 cm {x 9.5 cm; N 
31). In the turf (site 2), nests were 1-4 m 
apart. On March 5, 1992, the aggregation 
measured 2 X 25 m; there were 100 nests/ 
m- at site 1 and 30 nests/m- at site 2. The 
aggregation had about 2,900 nests in 1992. 
By March 1997, the 2 small pine trees near 
the aggregation had grown, shading it. the 
bees had moved most nests 2-3 m to the 
west, where it was sunnier. The winters and 
early springs of 1997 and 1998 were un- 
usually rainy. Fewer than 100 nests of A. 
fenningeri remained at site 1 by late March, 
1998 (1 nest/m- maximum density). 

Andrena fenningeri nested in the dense 
aggregation in only one small part of the 
large field (Fig. 2, Sites 1 and 2). Aggre- 
gated nesting by solitary bees and wasps is 
common, and some aggregations may per- 
sist at a site for over 50 years. There are 
several, nonexclusive explanations for this 
phenomenon, including substrate limitation; 
improved efficiency in foraging; protection 
from predators and parasites; attraction to 
others of the same species; an opportunity 
to save time and energy by re-using existing 
nests; and philopatry, or re-nesting near the 
insects' natal nests (see review in Rosen- 
heim 1990). Because A. fenningeri was ac- 
tive during early spring when the weather 
was often cold and rainy, the location of its 
aggregation in the warmest and sunniest 
part of the field appeared to be most advan- 
tageous; this maximized the number of 
hours available to the bees for foraging and 
reproduction. Floral resources were abun- 



VOLUME 101, NUMBER 1 



111 




Fig. 2. Field where A. fenningeri nested (drawn from aerial photograph). It is 230 m wide, and the arrow 
points North. Regularly mowed areas of short turf lawn are white (if sunny), striped (if partly shaded) or hatched 
(if always shaded). Stippled areas indicate locations without nests, being boggy, or fenced and shaded by dense 
tall grass and shrubs. Temperatures were recorded at locations (Sites) 1-6, all in red marine clay. The main 
portion of the nest aggregation was in a sunny patch of bare soil at Site 1 . and it extended into turf-covered 
soil (Site 2). There were no nests below the short turf at Sites 3-6; Site 3 was shaded all day; Sites 4 and 5 
were shaded in afternoons and mornings respectively; and Site 6 was sunlit all day. 



dant, thus not a limitation. The aggregation 
may have begun decades ago, when a mat- 
ed founding female successfully nested 
there, and her descendants returned to the 
area for nesting (philopatry). Because a 
closely related, cold-adapted species, A. 
(Scrapteropsis) alleghaniensis, placed its 
aggregation of nests in a southeasterly-fac- 
ing, sandy slope where they received max- 
imum insolation and warmth in early spring 
(Batra 1990), A. fenningeri was suspected 
of doing likewise, even though its aggre- 
gation was in level clay soil rather than in 
a sandy slope. 

In order to test this hypothesis, I made a 



series of temperature measurements at four 
depths (2, 8, 15, and 30 cm) at six sites 
(Sites 1-6, Fig. 2), using bimetallic-dial- 
probe thermometers. Most of the nest ag- 
gregation was at Site 1 (in exposed soil); 
some nests were in Site 2 (soil covered with 
short turf). 

A split-plot analysis was used on this ex- 
periment. The six sites were specifically se- 
lected for their shading and vegetation cov- 
erage conditions, and are therefore defined 
as fixed factors in this analysis. The sites 
are also defined as whole plots of the split- 
plot design, while the four levels of depth 
(2, 8, 15, 30 cm) at each site are the sub- 



112 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Table 1. Means and Standard Errors for the factorial combination of 4 depths and 6 sites. 



Depth 


2 




^ 




15 


30 




Site 


Mean*" 


SHM 


Meaii" 


SHM 


Mean'' 


SHM 


Mean" 


SHM 


Mean' 


1 


11.8 


0.77 


8.8 


0.65 


6.7 


0.63 


5.7 


0.56 


8.3^ 


1 


1 0.0 


0.88 


8.7 


0.73 


8.3 


0.71 


7.4* 


0.62 


8.6^ 


3 


2.5** 


0.82 


2 7** 


0.68 


2 6** 


0.66 


2.7** 


0.59 


2.6^ 


4 


8.1** 


0.89 


6.7* 


0.73 


6.7 


0.71 


6.1 


0.63 


6.9^ 


5 


8.2** 


0.89 


6.9* 


0.73 


6.1 


0.71 


5.1 


0.63 


6.6^ 


6 


9.9 

8.4^ 


0.88 


7.8 
6.9^^ 


0.73 


7.0 
6.2'' 


0.71 


6.7 
5.6'' 


0.62 


7.8^ 



■■ A least-squares-mean test examined possible difference between Site-1 (the one with the greatest nest con- 
centration) means and those of the other five sites. Within a column, a least-squares mean with one asterisk is 
significantly different from Site I's mean at 0.01 < P > 0.05. A mean with two asterisks is significantly different 
from Site I's mean at P < 0.01. 

^ Least-square mean. 

" Mean of all temperature measurements for each site for all four depths. 

'^ Mean of all temperature measurements for each depth for all six sites. 



plot levels. Measurements of temperature 
done on different days were defined as hap- 
hazard (a random number table was not 
used). Data were recorded during early af- 
ternoon on 7 days from March 3 to March 
26, 1992, during the time of peak nesting 
activity, and used as replication in the anal- 
ysis. A mixed-model analysis of variance 
was used to determine the fixed effects of 
site, depth, and their interaction. A hetero- 
geneous first-order autoregressive covari- 
ance structure was included in the ANOVA 
model to account for the possible associa- 
tions among depths. Least-significance dif- 
ference tests id = 0.05) were used to com- 
pare means of the fixed effects. Results are 
summarized in Table 1. 

Site 1 (the aggregation) was significantly 
warmer than site 3 at all depths (Table 1). 
It was warmer than Sites 4 and 5 at depths 
of 2 and 8 cm, but did not differ from them 
at 15 and 30 cm. Site 1 did not differ from 
Sites 2 and 6 at depths of 2, 8 and 15 cm, 
but, at 30 cm, it was slightly cooler than 
Site 2, and did not differ from Site 6. The 
right column of the Table shows the means 
of all measurements combined from the 4 
depths at each site. It shows that the portion 
of the aggregation at Site 2 was slightly 
(but not significantly) warmer than the ma- 
jor portion in Site 1, and both of these parts 
of the aggregation were considerably warm- 



er than areas where the bees did not nest, 
especially Site 3. 

The soil-temperature analysis indicates 
that A. fenningeri used the warmest local 
area for the aggregation. Thus, the bees 
thermoregulated by choosing a warm nest- 
ing site (and also individually, by basking). 
Sites 1 and 2 are at the southern edge of a 
pine-oak forest. The level ground receives 
maximum insolation at such a location in 
March (Kimball and Hand 1922, Geiger 
1965). The aggregation also benefitted from 
radiation that was reflected to it from near- 
by oak tree trunks (see Geiger 1965). Pine 
trees within the forest broke the cold north- 
west wind that prevailed on clear, sunny 
days in Maryland, and they retained heat 
that was radiated from the soil at night. The 
protective influence of a forest windbreak 
to the north of a field edge can raise air 
temperatures 5 cm above the ground all 
year, by 1 1 °C above air temperatures in a 
field near the forest along the opposite, 
south edge (Wales 1972). Such microcli- 
matic differences are significant; for ex- 
ample, 6 species of vernal wild flowers on 
a south-facing slope bloomed on average 6 
days earlier than the same species growing 
on a north-facing slope 50 m away; this dif- 
ference, correlated with cumulative differ- 
ences in air and soil temperatures, is equiv- 
alent to 176 km in latitude (Jackson 1966). 



VOLUME 101. NUMBER 1 



113 



There were slight differences within the 
aggregation between Sites 1 (exposed soil 
surface) and 2 (turf covered) that were not 
statistically significant, but could be detect- 
ed behaviorally when the temperature in the 
upper 2 cm of soil was near 1 1°C (marginal 
for flight initiation). For example, at noon 
on March 17, 1992, a sunny day after a 
frosty night, no bees were flying at either 
Site 1 or Site 2. At Site 1, temperatures 
were 1 1°C at 2 cm, 6.5°C at 8 cm, 4.5°C at 
15 cm, and 4.0°C at 30 cm depth. At 15: 
30, bees were flying at Site 1 but were not 
flying at Site 2. At Site 1, soil temperatures 
by then were 13°C at 2 cm, 10.5°C at 8 cm, 
8.0°C at 15 cm and 5.5°C at 30 cm depth. 
Temperatures then at Site 2 were too cool 
for flight, being only 10.5°C at 2 cm, 10.0°C 
at 8 cm, 9.5°C at 15 cm and 6.5°C at 30 cm 
depth. Although Site 2 was warmer than 
Site 1 at 15 and 30 cm, it was cooler at 8 
cm, and much (2.5°C) cooler at 2 cm depth, 
where bees usually wait to warm up, before 
flying. A similar pattern was seen at 13:00 
on March 25, 1992, a day of hazy sun after 
a frosty night: bees were flying at Site 1 but 
not at Site 2 (Site 1: 14.5°C at 2 cm, 9.0°C 
at 8 cm, 6.0°C at 15 cm, and 5.5°C at 30 
cm depth; Site 2: 1 1.0°C at 2 cm, 10.5°C at 
8 cm, 8.5°C at 15 cm, and 7.5°C at 30 cm). 
Thus, the sun warmed the upper layers of 
bare soil at Site 1 more quickly than the 
same depths in turf-shaded Site 2, permit- 
ting flight from Site 1 on cool, sunny days. 
The insulating turf retained warmth over- 
night in the lower layers at Site 2, but this 
did not promote flight activity. Even such 
slight differences between sites in bare soil 
and those in turf may be significant for sur- 
vival among bees that must fly to forage 
during inclement weather. 

Andrena nycthemera also begins adult 
activity when the soil thaws (as early as 
February 23 in 1990). Many more bees 
nested in a sunny part of the aggregation 
than in a shaded portion, where the soil re- 
mained frozen longer, and bees in sunlit ar- 
eas began seasonal activity several days be- 
fore those that nested in shady portions of 



the aggregation areas (Schonitzer and 
Klinksik 1990). Slight microclimatic differ- 
ences also influence the nesting behvior of 
halictine bees (Batra 1997, Potts and Will- 
mer 1997). 

Thermoregulation by Individuals 

Species of Andrena that were investigat- 
ed by Stone and Willmer (1989) produce 
relatively little endothermic heat, compared 
to some other genera of bees; instead they 
depend on insolation and warmth from sub- 
strates to generate the minimum 8-12°C 
ambient temperature needed to begin flight. 
Many species of Andrena bask at nest en- 
trances and on vegetation before takeoff on 
sunny, cool days. They include A. fulva 
Miiller (Paxton 1991), A. erigeniae Robt., 
which basks on fallen leaves that are 6-9°C 
warmer than ambient temperatures (Bar- 
rows 1978), and A. nycthemera, which can 
begin flights after basking at 8-10°C on 
sunny days, but cannot fly on cloudy days 
until the ambient temperature reaches 15°C 
(Schonitzer and Klinksik 1990). Herrera 
(1995) found that A. bicolor F. foraged on 
sunny days with an air temperature of at 
least 12-13°C; these small bees bask in 
warm microclimates to achieve the mini- 
mum internal thoracic temperature of 22°C 
needed to begin flight. 

Both sexes of A. fenningeri were able to 
begin flight on sunny days when the am- 
bient temperature of their microclimate was 
at least 1 1°C. Bees were often seen basking 
just inside nest entrances, on tumuli, be- 
neath or on fallen leaves, and on vegetation. 
The nearly black bodies of both sexes of A. 
fenningeri bear pale hairs, which are dens- 
est over the thorax (where warm-up of 
flight muscles is needed). In this way, the 
bees resemble the "heat-trap" structures of 
pussy willow flowers, which have been 
shown by Krog (1955) to absorb short- 
wave light (solar radiation), which passes 
through pale hairs into their black surfaces, 
and there becomes long wave radiation 
(heat), which is trapped in the dead airspace 
among the pale hairs. 



114 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Although A. fenningeri required a mini- 
mum, insolated temperature of 11°C to be- 
gin flying, they were active inside their 
nests at much lower temperatures. On Jan- 
uary 11, 1990, the soil at the aggregation 
(Site 1) had thawed after an unusually cold 
December 1989 (-20°C air temperature for 
several days, freezing the soil). I excavated 
twenty-seven cells on January 1 1 , to study 
the phenology of the bees and determine 
how they can emerge so early each spring. 
The air temperature (at 1 m, in shade) was 
10°C, and soil temperatures were 6°C at 2 
cm, and 4°C at 10, 21 and 30 cm depths. 
Some bees had already emerged from their 
cells (4 6 and 3 9 ); 9 males and 11 fe- 
males were still in their cells (sex ratio near 
1:1). Those in cells were resting on their 
backs. Bees that had emerged stood upright 
and had dug as far as 2 cm toward the sur- 
face; the earth that they had excavated was 
pushed backward, and packed into their na- 
tal brood cells, covering the fecal layer that 
had been deposited before their transfor- 
mation to prepupae late the previous May. 
When these cold (4°C) bees were disturbed, 
they vibrated their wings; when they were 
warmed slightly, they stood upright and be- 
gan to walk. These bees were stored in tis- 
sue culture wells at 4°C for 4 days. Some 
of them defecated (pale meconium), others 
chewed on moist soil that was placed in the 
wells with them. Thus, they were active at 
low temperatures. 

At the aggregation, air temperatures were 
above normal for January 1990, and by the 
18th, the air temperature at 14:00 E.S.T. (1 
m, shade) was 16°C, and soil temperatures 
were 13°C at 2 cm, 12°C at 10 cm (both 
above flight threshold), 10°C at 21 cm, and 
8°C at 30 cm. During January, the bees 
worked their way toward the surface, and 
the first bees (males) were flying on Feb- 
ruary 9, a sunny day, when the air temper- 
ature at 14:00 (at 1 m, in shade) was 17°C, 
and soil temperatures at Site 1 were 1 1 .5°C 
at 2 cm, 10°C at 10 cm, 8.5°C at 21 cm and 
7.5°C at 30 cm. Acer rubrum, Draba and 
Salix had just begun to bloom (Fig. 3). 



These early males flew slowly, about 2- 
7 cm above the aggregation (Site 1 only), 
frequently stopping to bask on dead oak 
leaves on the ground. Flying males dropped 
to the ground when clouds obscured the sun 
and when disturbed by the observer; they 
could then be captured by hand. On Feb- 
ruary 12, more nests were examined. One 
female had partly emerged from her cell at 
a depth of 17 cm, 3 others had emerged, 
and were in the soil at depths of 3, 9, and 
1 1 cm, having moved toward the surface. 
No males remained in their cells; 2 were 
found in the soil at 3 and 6 cm depths. Soil 
temperatures were 11°C at 2 cm, 8°C at 20 
cm and 6°C at 20 and 30 cm. Only males 
were seen again the next afternoon (sunny, 
15°C), circling low over the aggregation in 
calm air, but they settled on the ground and 
crawled around when gusts of wind or 
clouds arrived. The males made intermittent 
circular flights over small areas (20-30 cm 
radius), which gradually moved across the 
aggregation. No territoriality or male ag- 
gression was seen, and the males were not 
seen in nearby trees, where they formed 
mating swarms later in February. Female A. 
fenningeri released a pungent citrus odor 
from their mandibular glands when han- 
dled, but males had little odor. Possibly the 
patrolling males can smell the mandibular 
gland pheromone, released as the females 
dig their way toward the surface. 

On February 28, 1991, male behavior 
was similar. Fresh tumuli indicated that 
some females had begun nesting, but none 
were flying. Many males were patrolling 
near the ground in a zigzag pattern, often 
entering and leaving nests. When shaded, 
they dropped to the ground, and could be 
picked up by hand; they could not fly even 
though they buzzed vigorously, an activity 
that should warm their flight muscles. They 
were able to resume flight when replaced in 
the sun for about a minute. These small 
males could warm up in the sun at marginal 
temperatures more quickly than could the 
larger females, which may explain the ab- 
sence of flying females. The air temperature 



VOLUME 101, NUMBER 1 



115 



H S 



[-30 C 




rip- 4 months 



Fig. 3. Temperatures at the nest aggregation (Site 1) and phenology of A. fenningeri from January 1 1 through 
May 1, 1990, a warmer than nornial spring. A, air temperatures at 1 m height in shade; U, underground soil 
temperatures at a depth of 30 cm. Bees began emerging from brood cells in January; then crawled up the tunnels 
of their natal nests in early February. Red maples (R bar) bloomed from February 9 to March 21. The first bees 
(males) emerged February 9; female bees began collecting pollen from red maple by February 22; when maples 
finished blooming, bee activity declined and females switched to other pollen sources, including fruit trees (F). 
By May 4, the forest canopy had leafed out, the clay soil at the aggregation had dried and hardened, and only 
a few senile bees remained. From March 6-15, a record heat wave, with southwest winds, prevailed (H), and 
the bees were unusually active; on March 16, a cold front brought rain, and snow (S) fell on March 20 and on 
March 24-25. 



(14:30, in shade at 1 m) was 13.5-14.0°C; 
the soil (Site 1) at 2 cm was 10°C; at 20 
cm, 7.5°C and at 30 cm, 7°C. Males that 
had dropped to the ground to avoid wind 
often crawled beneath sunlit, dead, dry oak 
leaves, where temperatures were 12-15°C. 
In early March, in bright sun, temperatures 
under sunlit oak leaves reached 22°C when 
the ambient air temperature was 18°C. In 
early March 1992, some males that had 
been flying among trees in the mating 
swarm basked in the sun on dead leaves, 
but they crawled under the leaves when the 



wind gusted or clouds passed; some slept 
overnight in nest tunnels. When the air tem- 
perature at 1 m (shade) on a hazy day was 
at the 11°C threshold for flight, no female 
bees flew, but males were able to fly slowly 
for short distances between bouts of bask- 
ing in the weak sunlight. On a cloudy day, 
with air at 1 1°C, the temperature was only 
10°C below a fallen leaf, and also at 2 cm 
in the soil; insolation was insufficient on 
cloudy days, thus, no bees flew. 

Because A. fenningeri made more nests 
in the portion of the aggregation that was 



116 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



in bare soil (Site 1) than in the part that was 
in short (2-4 cm) turf (Site 2), and because 
bees emerged earher each spring at Site 1 
than at Site 2, soil temperatures at Site 1 
were compared with those at Site 2. Al- 
though no statistically significant tempera- 
ture difference between Sites 1 and 2 was 
detected in March, 1992 (see Table 1), there 
were small differences between the sites 
that may have been sufficient to account for 
the slightly earlier, and slightly more, activ- 
ity at Site 1, especially early in the spring, 
when temperatures marginally permitted 
flight. 

Temperatures were measured at Sites 1 
and 2 at the same time of day (early after- 
noon) on 13 days in March and April, 1990 
and 1993. Site 2 was cooler than Site 1 by 
a mean of 2.6°C at 2 cm, and by 1.1°C at 
8 cm; Site 2 was warmer than Site 1 by 
0.4°C at 15 cm and by 1.4°C at 30 cm 
depth. Evidently, the upper levels of bare 
soil (Site 1 ) warm more rapidly each sunny 
day than the same levels under grass, but 
the insulating grass (Site 2) retains warmth 
at lower levels overnight. The difference 
between Sites 1 and 2 was most prominent 
during sunny weather, for example on 
March 8, 1990, a sunny day after a clear, 
cold night, at 2 cm depth (where bees wait 
before flights). Site 1 was 14°C (warm 
enough for bees to fly), but Site 2 at 2 cm 
depth was only 10°C, which was below the 
flight threshold. However, after a several 
days of rain on a cloudy day, there was little 
difference between Sites 1 and 2. 

The behavior of male A. fenningeri in 
mating swarms was also influenced by mi- 
croclimate. At first, males flew only a few 
centimeters above the warm ground, where 
most nests were aggregated. As the air tem- 
perature warmed, many males moved their 
patrolling to the tips of the branches of 
small (4-5-m-tall) pine trees near the ag- 
gregation, where they formed swarms. On 
February 28, 1992, males swarmed around 
the pine trees at an air temperature (1 m, 
shade) of 17.5°C. When clouds or haze ar- 
rived, the males suddenly ceased flying. 



alighted, and crawled between the dark 
green pine needles (which would retain 
heat); they resumed their flights when the 
sun reappeared. 

Behavior of Males 

The behavior of male Andrena bees has 
been described and reviewed by Barrows 
(1978), Gebbardt and Rohr (1987), and 
Hallmen (1991). Males may emerge simul- 
taneously with, or earlier than, their females 
each year. They swarm conspicuously in the 
sunshine, circling and zigzagging above ag- 
gregations of nests, around flowers of host 
plants, and around tall landmarks, such as 
selected trees near aggregations. Males of 
most species are non-territorial, and they 
jointly patrol in search of females, without 
noticeable interactions among males. Males 
of A. nycthemera are unusually aggressive, 
biting each other and competitively digging 
in the ground, searching for emerging fe- 
males (Shonitzer and Klinksik 1990). De- 
spite the large numbers of both sexes that 
emerge and mate within a few days, and the 
numerous times males are seen pouncing on 
females and on various small dark objects, 
actual copulation is surprisingly rarely ob- 
served. I also did not observe the mating of 
A. fenningeri, in spite of many hours spent 
watching their behavior. 

Both sexes of A. fenningeri produce a 
lemon-like odor from their mandibular 
glands when captured. It is most distinct in 
females. The mandibular gland secretions 
of some Andrena bees contain complex 
mixtures of spiroacetals, monoterpenes, and 
other compounds (Bergstrom and Tengo 
1982). The secretion of A. fenningeri prob- 
ably includes geraniol and citronellol, 
which are major components in other spe- 
cies of Andrena (Bergstrom and Tengo 
1982). Male bees use these secretions to 
mark the areas that they patrol in search of 
females (Bergstrom and Tengo 1982, Geb- 
hardt and Rohr 1987, Hallmen 1991); and 
female bees use both mandibular-gland and 
Dufour's-gland secretions to mark and iden- 



VOLUME 101. NUMBER 1 



117 



tify their nests (Ayasse et al. 1990; Stein- 
mann 1990). 

During an unusually warm spring, the 
first male A. fenningeri emerged on Febru- 
ary 9, 1990, before the females. They 
emerged as late as March 7, in 1993, also 
before the females appeared. Some years, 
males flew before any host plants bloomed; 
in other years, the first bees appeared when 
hosts started blooming. Andrena nycthem- 
era males also may begin flying before any 
food is available (Schonitzer and Klinksik 
1990). During their first activity, males 
crawled, or patrolled, flying in the sunshine 
some 2-8 cm above the bare soil at Site 1, 
which radiated heat, warming the relatively 
calm layer of air near the ground. They flew 
upwind in small circles or in slow, waver- 
ing patterns, occasionally waiting on fallen 
leaves when clouds passed. They also fell 
to the ground and "froze" when alarmed 
(by my movements) and could be caught 
with the fingers. On cool days, newly 
emerged females also displayed such than- 
atosis, when disturbed, becoming immobile, 
dropping to the ground, with legs held stiff- 
ly, parallel and close to their bodies, a be- 
havior similar to that of elaterid beetles. 
Sometimes, such bees relaxed, and slowly 
crawled beneath dead leaves. This behavior 
was seen only in early spring, before host 
plants began to bloom. Perhaps it is an en- 
ergy-conserving defense mechanism, used 
at a time when the bees were subsisting on 
their stores of fat. Male A. fenningeri were 
seen entering and leaving exit holes and 
nest entrances. Some males slept in holes, 
but they did not dig for females. Occasion- 
ally, both males and females crawled on the 
surface of the aggregation, but mating was 
not seen there. 

Most females emerged each year after 
one or more warm, sunny days, and fresh 
tumuli indicated that nesting had begun. At 
this time, many male A. fenningeri ceased 
patrolling at the aggregation, and instead, 
they swarmed among the bare branches of 
an oak tree and around two small pines {P. 
virginiana Mill.) growing next to the ag- 



gregation. Males of both Nomada sayi 
Robt. and N. perplexa Cress, swarmed to- 
gether with the males of A. fenningeri, both 
at the aggregation and around the trees; per- 
haps they have similar pheromones. Male 
A. fenningeri were seen to emerge from the 
ground and fly up to join these others 
around the trees. 

Males of A. fenningeri patrolled areas 
around the tips of the pine branches. They 
zigzagged along the downwind sides of the 
branches at a height of 2-4 m, occasionally 
alighting. They flew from one branch tip to 
the next, briefly hovering while facing the 
tip of each branch, before flying upwind to 
the next branch, where they zigzagged to- 
ward its tip. This was repeated, until they 
reached the upwind (and sunny) side of the 
tree, from which they drifted back on the 
breeze, to repeat the process of inspecting 
branch tips. Mating was not observed at 
these "swarm trees." Similar upwind and 
zigzag patrolling, alternating with down- 
wind drifting, occurs in A. vaga (Hallmen 
1991). 

On March 2, 1992, an attempt to attract 
males to females in a swarm was made. 
Four live females were tied with threads to 
the tips of pine branches where many males 
patrolled. They were conspicuous, but were 
bypassed by 16 males. Another 7 males 
hovered to briefly inspect them, and 4 
males pounced on the females, but imme- 
diately released without mating. These fe- 
males orally released a distinct lemon-like 
odor. Other bees that were swarming 
around the pine trees were netted, yielding 
10 males and 1 female, indicating that fe- 
males were also attracted to these "swarm 
trees" (see Hallmen 1991). The captured 
female and 5 males were dissected; the fe- 
male was inseminated and had fed on maple 
pollen, but the males had not eaten pollen 
or nectar, their crops being collapsed and 
their guts empty. A sample of 6 females 
that were flying over the aggregation on 
March 2 had all eaten maple pollen, and all 
were inseminated. Similarly, on February 
28, 1992, 14 males that were swarming 



118 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



around the pine trees were dissected; 1 2 had 
empty crops and guts, 1 had eaten nectar, 1 
had eaten some maple pollen; and most had 
large fat bodies. The females that were col- 
lected from nests with tumuli on that day 
varied: 1 had not mated or eaten, and still 
had rectal meconium; 2 were inseminated 
but had not eaten. The number of males 
seen around the swarm trees and above the 
aggregation each year slowly declined as 
spring advanced, and a week to 10 days af- 
ter emergence, males were no longer seen 
near the aggregation. By this time, host 
plants were blooming, and the males 
searched for females there. 

Parasites and Associates 

As with other species of And ren a world- 
wide, A. fenningeri was associated with 
several other insects at the aggregation. 
These included the halictine bee, Dialictus 
versatus (Robertson), nesting at the western 
edge of the aggregation, and its cleptopar- 
asite, Sphecodes sty gins Robertson. Five 
species of Nomada were active at the A. 
fenningeri aggregation, which I identified 
as N. bella Cresson, N. cressoni Robertson, 
N. parx'a Robertson, N. perplexa Cresson 
and N. sayi Robertson. Nomada perplexa 
was by far the most abundant of these clep- 
toparasitic anthophorid bees. They often en- 
tered nests, but I did not find any Nomada 
in brood cells. A conopid fly (probably My- 
opa sp.), bombyliid flies, blue and black 
fungi in brood cells, and three species in 
the genus Eustalomyia (Anthomyiidae) also 
were present. These anthomyiids were at- 
tracted to flying A. fenningeri, but not to D. 
versatus, which had a different, more erratic 
flight pattern. 

The grouping of nests in a perennial ag- 
gregation permitted a permanent population 
of these natural enemies, but the bees had 
several defenses against them. For example, 
many nests were initiated beneath dead 
leaves, which concealed their entrances; tu- 
muli often were not rebuilt when they had 
been destroyed by rain, which made nest 
entrances inconspicuous; nest entrances 



were temporarily sealed with soil particles; 
returning foragers dodged cleptoparasites 
that followed them; foragers took circuitous 
routes, and entered nests abruptly, thus con- 
fusing and evading trailing parasitic bees 
and flies. Such evasive techniques have 
been seen in other species of solitary bees 
and wasps (Hager and Kurczewski 1985, 
Meyer-Holzapfel 1986, Rosenheim 1990, 
Schonitzer and Klinksik 1990). 

The most abundant of all the parasites 
were three Eustalomyia species (det. W. 
Downes, Jr.). The only hosts previously 
known for this genus of anthomyiid flies are 
the solitary crabronine wasps, Ectemnius 
paucimaculatus (Packard) in North Ameri- 
ca (Krombein 1964), and E. cavifrons 
(Thomson) in Europe (Meyer-Holzapfel 
1986). According to Downes (in litt.), the 
three species that are associated with A. fen- 
ningeri may be the first to be associated 
with bees, and probably are new records, 
because these flies appeared much earlier in 
spring than those species that are associated 
with crabronine wasps. 

The Eustalomyia species appeared in 
succession; of samples sent for identifica- 
tion, the first (species A, males) being ac- 
tive at the aggregation as early as March 9, 
1992, 12 days after the first bees emerged 
that year. Females of species A were active 
at the aggregation March 15-22, 1990, and 
March 25, 1988; species-A males were 
found as late as April 27, 1988. Females of 
species B were collected at the aggregation 
on April 4, 1988, and females of species C 
were present on April, 27, 1988. 

The behavior and gross morphology of 
all three species of Eustalomyia were sim- 
ilar. It was not possible to distinguish 
among them in the field; thus, they will be 
discussed collectively here. Initially, they 
were assumed to be the anthomyiids, Leu- 
cophora obtusa (Zetterstedt), and L. mary- 
landica (Malloch), which are common at 
aggregations of three species of Colletes 
bees at Beltsville, and behave similarly to 
Eustalomyia. Leucophora obtusa also par- 



VOLUME 101, NUMBER 1 



119 



asitizes Andrena nycthemera in Europe 
(Schonitzer and Klinksik 1990). 

Cleptoparasitic anthomyiid and sarcoph- 
agid flies are often classified according to 
their behavior in relation to their hosts. 
Some are "satellite flies," also called "sta- 
tion takers," which perch near the nests of 
hosts, abruptly taking flight to pursue re- 
turning foragers to their nests. Others are 
"hole searchers," flying about in search of 
hosts' nests (Hager and Kurczewski 1985, 
Wcislo 1986, Meyer-Holzapfel 1986). The 
host bees and wasps defend their nests 
against satellite flies by evasive maneuvers 
in flight. Intense activity by both hosts and 
parasites at dense aggregations also causes 
confusion of the parasites. Closure and con- 
cealment of nest entrances provide some 
protection from hole-searching flies and the 
cleptoparasitic bees, Nomada and Spheco- 
des (see review in Rosenheim 1990). 

A 1-m- area in the densest part of the 
aggregation (54 nests/m-^ on March 8) was 
watched for 74 minutes on a warm, sunny 
day (10:10-11:24, March 15, 1990), in or- 
der to observe the activities of cleptopar- 
asites, especially Eiistalomyia. During this 
time, A. fenningeri foragers left nests 4 
times and returned with yellow pollen 10 
times (one trip lasted 14 minutes); two bees 
remained at nest entrances; usually, one A^o- 
mada parva continually patrolled above the 
nests; once one alighted briefly, to fan her 
wings at a nest entrance (probably to bring 
up odors from the nest). 

Most of the time, two EustaJomyia were 
perched in the area. The flies jumped up 
and briefly followed patrolling Nomada 
twice; they chased each other once; and one 
briefly followed a falling leaflet. One or two 
flies 10 times ioWowQd A. fenningeri, while 
both leaving and returning to their nests. 
Flies were seen entering A. fenningeri nests 
3 times. In one instance, two flies followed 
a bee returning with pollen; one of them 
alighted and peered into the nest headfirst, 
then turned and slowly backed into the nest, 
until it disappeared. After 4 minutes, it re- 
appeared at the entrance, waited 10 sec- 



onds, and flew away (the bee remained in- 
side). Another time, two flies followed a 
bee returning with pollen (she zigzagged in 
an attempt to lose them); one fly followed 
this bee into her nest headfirst, but left with- 
in a minute. Once, two flies followed a bee 
laden with pollen to her nest entrance, 
which was beneath a fallen leaf. On alight- 
ing near the nest, the flies shoved each other 
for 1-2 seconds, until one of them flew 
away. The "winner" then slowly backed 
tailfirst into the nest entrance, rested there 
for 2 minutes, then backed down into the 
tunnel, out of sight. After another 4 min- 
utes, the fly's head reappeared at the en- 
trance, the fly walked out of the nest and 
rested for 2 minutes facing away from it, 
cleaned itself, and departed. 

The bees defended their nests in several 
ways. The entrances to many nests were 
closed with plugs of soil or obscured by 
loose tumuli. Entrances to other nests were 
camouflaged by, or hidden beneath, dead 
leaves, grass blades, and twigs. Bees that 
appeared to be ready to leave their nest en- 
trances on foraging trips were hesitant, 
backing down when other insects flew near- 
by; departing bees were briefly followed by 
flies, but they did not seem to take evasive 
paths. Ten bees, returning to their nests that 
were followed by Eustalomyia, performed 
elaborate evasive maneuvers before abrupt- 
ly diving into their nests' entrances. In con- 
trast, two returning bees that were not fol- 
lowed by flies entered their nests directly. 
During evasive flights, returning bees pur- 
sued by flies zigzagged; one of them left 
the area for 2 minutes, returning to her nest 
again without the fly; another bee shook off 
two closely-following flies when she 
crawled beneath a leaf to the hidden en- 
trance of her nest; a third bee had nested in 
the shadow of a large piece of debris; the 
pursuing fly did not follow her, but alighted 
in a sunny area nearby. Another bee had 
nested in turf; this bee hit the grass blades 
and fell to the ground when she tried to 
zigzag to evade two pursuing flies. The 
placement of nests in shadows beneath de- 



120 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



bris may help prevent Eustalomyia attacks, 
but this may also be disadvantageous be- 
cause less warmth (insolation) would be 
present at such shady sites. 

Adult Eustalomyia spp. first appreared at 
the aggregation soon after the bees began 
to emerge. Nomada perplexa emerged at 
the same time as their hosts, and even 
shared mating aggregation sites with A. /<?/?- 
ningeri. On cool, sunny days in early 
spring, Eustalomyia, Nomada, and their 
hosts basked on the warm soil at the aggre- 
gation. The initiation of adult activity by 
these insects in February through March 
varied by up to a month in different years, 
but the time of its termination each year in 
early May varied by only about a week. 

Only one small maggot was found (May 
4, 1989), although 22 nests of A. fenningeh 
were examined, and Eustalomyia were 
abundant. It was on the side wall of a sealed 
cell that contained a normal egg on an in- 
tact pollen ball, in a marked nest. This nest 
was closed at the surface, with only this one 
cell, at the end of a backfilled lateral. This 
cell was being provisioned on April 21, and 
Eustalomyia had been very active at the ag- 
gregation. At that time, this nest had a tu- 
mulus; probably it was a new, secondary 
nest. There was no bee in this closed nest 
when it was examined on May 4. The genus 
Eustalomyia is ovoviviparous (Meyer-Hol- 
zapfel 1986). Those species associated with 
crabronine wasps oviposit at the host's nest 
entrances; the fly larvae hatch immediately, 
and crawl to the wasps' food stores (Meyer- 
Holzapfel 1986). In contrast, the species 
that are associated with A. femiingeri enter 
the bee's nests, even while the adult bee is 
still inside, and they somehow manage to 
deposit their eggs (or perhaps larvae) inside 
cells that have been completed, but not yet 
sealed (the cell plug and long, backfilled 
lateral would probably prevent these deli- 
cate flies and their maggots from entering 
sealed cells). Because solitary bees imme- 
diately begin to seal their brood cells and 
backfill laterals on completing oviposition, 
which occurs after a period spent smooth- 



ing and perfecting the pollen ball (Batra 
1970), it would be difficult for the flies to 
seize this opportunity. If flies oviposited in 
cells before pollen balls are completed, the 
gyrations and grooming of the pollen ball 
by the bees working in the confinement of 
their cells would damage fly eggs or larvae, 
and if detected, the bees may kill, eat, or 
remove them. On March 22, 1990, one Eu- 
stalomyia fly that had backed into a nest 
that was occupied by a bee, as if to oviposit, 
rapidly emerged while buzzing its wings, 
agitated, as if it had been attacked by the 
defensive bee. It then waited, 1 mm from 
the nest entrance and facing it, for the next 
33 minutes. Perhaps the flies deposit their 
eggs in crevices in open laterals and the 
hatchling larvae slip into the open cells 
while the bees are temporarily quiescent, 
during the process of oviposition. 

Conopid flies also parasitize these bees. 
On May 7, 1987, two dead female A. fen- 
ningeri were found, one in each of two 
nests. In both instances, the bee, with swol- 
len abdomen, was poised over a completed 
pollen ball, as if to begin oviposition (Fig. 
ID). One of them contained a brown pu- 
parium; a large maggot emerged from the 
pulsating abdomen of the other bee 30 min- 
utes after her collection. Evidently, the 
pressure of the maggots inside the bees 
mimicked the stimulus of eggs that were 
ready to lay, causing the bees to prepare 
normal-appearing pollen balls, and take the 
position for egg laying, as they died (there 
may have been additional stimuli). Both 
nests had other cells with eggs on pollen 
balls, suggesting that these bees had laid 
eggs soon before the rapidly-growing mag- 
gots consumed most of their abdominal 
contents. According to Smith (1966), An- 
drena bees are usually parasitized by spe- 
cies of conopid flies in the genus Myopa; 
host bees may actively fly and feed with a 
large maggot nearly filling the abdomen; 
death of the bee occurs shortly before the 
maggot's pupation. 



VOLUME 101, NUMBER 1 



121 



Conclusion 

Andrena fenningeri is a species of soli- 
tary, vernal, univoltine bee that successfully 
exploits ecological resources that become 
available as soon as the ground thaws. In 
order to be prepared for the earliest bloom, 
this bee nests in permanent aggregations in 
the warmest available microhabitat. Over- 
wintered adults begin to move out of their 
natal brood cells toward the surface in mid- 
winter On emergence from nests, they 
mate, often before food is available, and 
soon females begin to excavate new nests 
in the aggregation. When the temperature is 
marginal for flight, these black bees bask in 
the sun, to gain sufficient warmth for their 
activities. 

This species first feeds on early-bloom- 
ing red maple and willow, but later shifts 
to pear and peach as they begin to bloom. 
It may be possible to manage this species 
for orchard pollination, if suitable nesting 
sites and early-season hosts are provided. 
Areas of sunny, exposed, level clay soil 
along the north edges of orchards could be 
prepared and kept free of vegetation. They 
should be backed along their north sides by 
windbreaks, such as forests, walls, or solid 
fences. Red maples and willows should 
grow within 100 m. The initial population 
of bees could be obtained by transplanting 
cores of soil that contain nests from an ex- 
isting aggregation, as is done to move soil- 
dwelling alkali bees (Batra 1970). Once a 
population of bees is established near the 
orchard, it would become permanent and 
maintenance-free, except for the need to re- 
move (scrape) all vegetation from it each 
winter to permit maximum insolation in 
early spring. 

Acknowledgments 

I thank Dr L. W. Douglass and P. Pena- 
fiel of the Statistical Consulting Services, 
USDA, Beltsville, for the statistical analy- 
ses of soil temperatures. Dr. W. E. La Berge 
of the Illinois Natural History Survey kind- 
ly identified A. fenningeri. Dr. W. L. 



Downes, Jn of Grand Rapids, Michigan, 
identified the Eustalomyia and provided in- 
formation about this uncommon genus of 
flies. Reviews by Drs. E. M. Barrows of 
Georgetown University and B. B. Norden 
of the National Museum of Natural History 
improved this publication. 

Literature Cited 

Ayasse, M., R. Leys, F Pamilo and J. Tengo 1990. 
Kinship in communally nesting Andrena (Hyme- 
noptera: Andrenidae) bees, indicated by the com- 
position of Dufour's gland secretion. Biochemical 
Systematics and Ecology 18: 453-460. 

Barrows, E. M. 1978. Male behavior in Andrena eri- 
geniae (Hymenoptera: Andrenidae) with compar- 
ative notes. Journal of the Kansas Entomological 
Society 51: 798-806. 

Batra, S. W. T. 1970. Behavior of the alkali bee, Nomia 
melanderi, within the nest. Annals of the Ento- 
mological Society of America 63: 400-406. 

. 1985. Red maple (Acer rubrum L.), an im- 
portant early spring food resource for honey bees 
and other insects. Journal of the Kansas Entomo- 
logical Society 58: 169-172. 

. 1990. Bionomics of a vernal solitary bee, A/i- 

drena (Scrapteropsis) alleghaniensis Viereck, in 
the Adirondacks of New York (Hymenoptera: An- 
drenidae). Journal of the Kansas Entomological 
Society 63: 260-266. 

. 1997. Bionomics of Lasioglossum (Evylaeiis) 



niatianense (Bliithgen) (Hymenoptera: Halicti- 
dae), the predominant pollinating bee in orchards 
at high altitude in the Great Himalaya of Garhwal, 
U.R India. Proceedings of the Entomological So- 
ciety of Washington 99: 162-170. 

Bergstrom, G. and J. Tengo. 1982. Multicomponent 
mandibular gland secretions in three species of 
Andrena bees (Hymenoptera, Apoidea). Zeit- 
schrift fiir Naturforschung 37c: 1 124-1 129. 

Chambers. V. H. 1968. Pollens collected by species of 
Andrena (Hymenoptera: Apidae). Proceedings of 
the Royal Entomological Society, London (A) 43: 
155-160. 

Gebhardt. M. and G. Rohr. 1987. Zur Bionomie der 
Sandbienen Andrena clarkella (Kirby), A. ciner- 
aria (L.), A. fuscipes (Kirby) und ihrer Kuckucks- 
bienen (Hymenoptera: Apoidea). Drosera 87: 89- 
114. 

Geiger, R. 1965. The Climate Near the Ground. Har- 
vard University Press, Cambridge. 611 pp. 

Hager, B. J. and F. Kurczewski. 1985. Cleptoparasitism 
of Ammophila harti (Fernald) (Hymenoptera: 
Sphecidae) by Senotainia vigilans Allen, with ob- 
servations on Phrosinella aurifacies Downes 
(Diptera: Sarcophagidae). Psyche 92: 451-462. 

Hallmen, M. 1991. Einige Beobachtungen zum Flug- 



122 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



verhalten von Drohnen an einer Kolonie der So- 
litarbiene Andrena vaga Panzer (Hymenoptera: 
Andrenidae). Nachrichten des Entomologischen 
Vereins Apollo, Frankfurt/Main N. F. 12: 107- 
120. 

Herrera, C. M. 1995. Floral biology, microclimate, and 
pollination by ectothermic bees in an early-bloom- 
ing herb. Ecology 76: 218-228. 

Hurd, P D., Jr 1979. Apoidea. pp. 1741-2209. //; 
Krombein, K. V., D. R. Smith, and B. D. Burks, 
eds.. Catalog of Hymenoptera in America North 
of Mexico, Vol. 2. Smithsonian Institution Press, 
Washington, D. C. 

Jackson, M. T 1966. Effects of microclimate on spring 
flowering phenology. Ecology 47: 407-415. 

Kimball, H. H. and I. E Hand. 1922. Daylight illumi- 
nation on horizontal, vertical, and sloping surfac- 
es. Monthly Weather Review 50: 615-627. 

Krog, J. 1955. Notes on temperature measurements in- 
dicative of special organization in Arctic and Sub- 
arctic plants for utilization of radiated heat from 
the Sun. Physiologia Plantarum 8: 836-839. 

Krombein, K. V. 1964. Natural history of Plummer's 
Island, Maryland. XVII. The hibiscus wasp, an 
abundant rarity, and its associates (Hymenoptera: 
Sphecidae). Proceedings of the Biological Society 
of Washington 77: 72-112. 

La Berge, W. E. 1971. A revision of the bees of the 
genus Andrena of the Western Hemisphere. Part 
IV. Scropteropsis, Xiphandrena and Rhaphan- 
drena. Transactions of the American Entomolog- 
ical Society 97: 441-520. 

Meyer-Holzapfel, M. 1986. Vergleichende Betrachtung 
des Verhaltens brutparasitischer Fliegen aus zwei 
Familien bei solitaren Aculeaten Hymenopteren 
(Methodische Kritik). Milleilungen der Schwei- 
zerischen Entomologischen Gesellschaft 59: 95- 
110. 

Miliczky, E. R. and E. A. Osgood. 1995. Bionomics 
of Andrena (Melandrena) vicina Smith in Maine 
and Washington, with new parasite records for A. 
(M.) regularis Malloch and a review of Melan- 
drena biology. Journal of the Kansas Entomolog- 
ical Society 68: 51-66. 



Paxton, R. J. 1991. Profile of a solitary bee: Andrena 
fulva. Bee World 72: 11-18. 

Potts, S. G. And P Willmer. 1997. Abiotic and biotic 
factors influencing nest-site selection by Halictiis 
rubicundiis, a ground-nesting halictine bee. Eco- 
logical Entomology 22: 319-328. 

Riddick, E. W. 1990. Andrena macra Mitchell (Hy- 
menoptera: Andrenidae) overwinter and delay 
spring emergence in Virginia. Proceedings of the 
Entomological Society of Washington 92: 771- 
772. 

. 1992. Nest distribution of the solitary bee An- 
drena macra Mitchell (Hymenoptera: Andreni- 
dae), with observations on nest structure. Pro- 
ceedings of the Entomological Society of Wash- 
ington 94: 568-575. 

Rosenheim, J. A. 1990. Density-dependent parasitism 
and the evolution of aggregated nesting in the sol- 
itary Hymenoptera. Annals of the Entomological 
Society of America 83: 277-286. 

Schonitzer, K. and C. Klinksik. 1990. The ethology of 
the solitary bee Andrena nycthemera Imhoff 1866 
(Hymenoptera. Apoidea). Entomofauna, Zeit- 
schrift fiir Entomologie 11: 377-427. 

. 1992. Individuell unterschiedlicher Leben- 

slauf bei der Sandbiene Andrena nycthemera Im- 
hoff. Nachr Blatt Bayerischen Entomologie 39: 
116-121. 

Smith, K. G. V. 1966. The larva of Thecophora occi- 
densis, with comments on the biology of Conopi- 
dae (Diptera). Journal of Zoology, London 149: 
263-276. 

Steinmann, E. 1990. Zur Nahorientierung der solitaren 
Sandbiene Andrena vaga. Panzer 1799 (Hyme- 
noptera. Apoidea) am Nesteingang. Bulletin de la 
Societe Entomologique Suisse 63: 77-80. 

Stone, G. N. and P G. Willmer. 1989. Warm-up rates 
and body temperatures in bees: the importance of 
body size, thermal regime and phylogeny. Journal 
of Experimental Biology 147: 303-328. 

Wales, B. A. 1972. Vegetation analysis of north and 
south edges of a mature oak-hickory forest. Eco- 
logical Monographs 42: 451-471. 

Wcislo, W T 1986. Host nest discrimination by a clep- 
toparasitic fly, Metopia campestris (Fallen) (Dip- 
tera: Sarcophagidae: Miltogramminae). Journal of 
the Kansas Entomological Society 59: 82-88. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999. pp. 123-137 

REVISION, CLADISTICS AND BIOGEOGRAPHY OF THE NEOTROPICAL 
GENUS SOUZALOPESMYIA ALBUQUERQUE (DIPTERA: MUSCIDAE) 

Claudio Jose Barros de Carvalho 

Department of Zoology, Universidade Federal do Parana, C.P 19020, 81531-990, Cu- 
ritiba, Parana, Brazil (e-mail: cjbcarva@bio.ufprbr) 



Abstract. — Souzalopesmyia Albuquerque, a monophyletic Neotropical muscid genus 
of five species, is reviewed to include two new species, Souzalopesmyia paraensis 
Carvalho, new species (Brazil: Para), and Souzalopesmyia sulina Carvalho, new spe- 
cies (Paraguay: Canindeyu). Ground plan characters of the Phaoniinae as outgroup were 
used in a cladistic analysis of the genus. The phylogenetic relationships found are (S. 
amazonica {S. paraensis (S. singularis, S. sulina)) (S. carioca)) and these seem to sup- 
port at least two biogeographical hypotheses: 1, The basal clade, S. amazonica, suggests 
a date of origin for the genus as far back as the Late Cretaceous; 2, The occurrence of 
5'. paraensis in Belem, along south side of the Amazon River, suggests a single dispersal 
event to colonize that region, in a more recent time, which belongs to the northwestern 
track. 

Resumo. — Souzalopesmyia Albuquerque, um genero monofiletico de Muscidae Neo- 
tropical com tres especies e revisto para incluir duas novas especies, Souzalopesmyia 
paraensis Carvalho (Brasil: Para) e Souzalopesmyia sulina Carvalho, (Paraguay: Can- 
indeyu). Para a analise cladistica do genero, foram utilizados, como grupo de fora, os 
caracteres do piano basico de Phaoniinae. A partir do relacionamento filogenetico encon- 
trado {S. amazonica (S. paraensis {S. singularis, S. sulina)) (S. carioca)) podem ser re- 
tiradas, no minimo, duas hipoteses biogeograficas: 1. O clado basal, S. amazonica, sugere 
a data de origem do genero para o Cretaceo Superior; 2. A ocorrencia de S. paraensis 
em Belem, ao sul do Rio Amazonas, sugere que um unico evento de dispersao ocorreu 
para colonizar esta regiao, em uma epoca mais recente. 

Key Words: Cladistics, biogeography, phylogenetic analysis, Souzalopesmyia, taxonomy 



Souzalopesmyia Albuquerque 1951 is an other genera of Muscidae is confusing, 

unusual genus of rare Neotropical Muscidae Malloch (1929) described Peruvia (a pre- 

because all of its species are yellow. In gen- occupied name, now a synonym of Souza- 

eral, yellow species are inhabitants of very lopesmyia) to include only Mydaea singu- 

dark shaded habitats. The genus was pro- laris Stein, and considered it to be close to 

posed by Albuquerque (1951: 53) to ac- Charadrella Wulp 1896 (Cyrtoneurininae). 

commodate two new species, S. carioca Seguy (1937) synonymized Peruvia with 

and S. amazonica. Subsequently, Pont Mydaea Robineau-Desvoidy 1830 (Myda- 

(1972) included Mydaea singularis Stein einae), and Albuquerque (1951) considered 

1911, in the genus. Souzalopesmyia to be close to Oramydaea 

The relationship of Souzalopesmyia with Snyder, the latter being an Afrotropical ge- 



124 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



nus of Mydaeinae now synonymized with 
Myospila Rondani 1856. Hennig (1965), af- 
ter examination of the ovipositor of the type 
species of Peruvia put it close to Helitia 
Robineau-Desvoidy (Phaoniinae). Carvalho 
(1989d) included Soiizalopesmyia in his 
cladistic analysis of Muscidae and consid- 
ered it as genus of Phaoniinae (Pont 1972, 
Carvalho et al. 1993). 

The present paper adds two new species 
to Soiizalopesmyia, presents a cladogram of 
the genus, and reflects upon the biogeo- 
graphic relationships among the included 
species. 

Material and Methods 

Specimens from the following institu- 
tions were studied for comparative purpos- 
es: Department of Zoology, Universidade 
Federal do Parana (DZUP), Curitiba, Bra- 
zil; Museu Paraense Emflio Goeldi 
(MPEG), Belem, Brazil; Museu Nacional 
do Rio de Janeiro (MNRJ), Rio de Janeiro, 
Brazil; Staatliches Museum fiir Tierkunde 
(SMT), Dresden, Germany; Museo Nacion- 
al de Historia Natural (MNPA), Asuncion, 
Paraguay. 

The terminology and abbreviations used 
here are those in McAlpine (1981) and Car- 
valho (1989a), and the descriptions of the 
type-specimen labels follows O'Hara 
(1982). 

The sister group of Soiizalopesmyia is 
unknown although the genus is considered 
as one of the most basal members of the 
tribe Phaoniini (Pont and Carvalho 1997). 
The sister group may be found in Afrotrop- 
ical Phaoniini (Pont 1980), which is com- 
posed of three genera. Two of these genera, 
Phaonia Robineau-Desvoidy, 1830 and 
Helina, are true but paraphyletic genera of 
Phaoniini (Hennig 1965). On the other 
hand, the sister group may be found in a 
more basal group of Neotropical Muscidae. 
Character polarities for the genus were 
based on the ground plan of Phaoniinae, 
and the assignment by Pont (1986) since no 
phylogenetic analysis for these genera is 



Table 1. Character state distribution among species 
of Soiizalopesmyia. = plesiomorphic character 
states; 1 = apomorphic character states; ? = missing 
data. Taxonomically useful characters for the species 
of Soiizalopesmyia. Characters with an * were used in 
phylogenetic analysis. 



outgroup 
S. amazonica 
S. carioca 
S. paraensis 
S. singularis 
S. sulina 



000 000 000 000 00 

000 100 010 101 11 

101 001 111 101 11 

000 oil 110 001 11 

000 oil 100 111 11 

?10 oil 100 111 11 



*5. 



*6. 

*7. 



10. 



12. 



13. 
14. 



Number of frontal setae in female; 0) three; 1) 
two. 

Frontal setae in male: 2) two cruciate; 1 ) two, the 
lower cruciate, the upper reclinate. 
Vti: 0) parallel; 1) divergent. 
Postocular setae row in male: 0) complete and 
distinct, whole row of setulae reaching epistome; 
1 ) incomplete, row of setulae reaching only to 
basal half of eyes. 

Postocular setae row in male: 0) whole row of 
setulae black; 1 ) composed of black and yellow 
setulae; latter beginning after basal half of eye 
and reaching epistome, but 1-2 with black setu- 
lae. 

Number of Dc prst: 0) two; 1) one. 
Acr female prst: 0) distinct from the ground se- 
tulae; 1 ) not distinct from the ground setulae. 
Proepisternal seta: 0) strong, similar in length to 
the upper anepistemal setae; 1 ) weak, less than 
the length of upper anepistemal. 
Crossvein dm-cu: 0) almost straight (Albuquer- 
que 1951: Fig. 11); 1) weakly curved (Albu- 
querque 1951: Fig. 13). 

Fifth sternite shape: 0) without sharp depression 
on posterior side (Fig. 1); 1) with sharp depres- 
sion on posterior side (Figs. 2, 3, and Albuquer- 
que 1951: Figs. 10 and 12). 

Cereal plate: 0) round outline (Fig. 4, and Al- 
buquerque 1951: Fig. 4); 1) squared outline 
(Figs. 5, 6). 

Head appearance: 0) not elongate; 1 ) elongate 
(Albuquerque 1951: Fig. 1). 
Number of ocellar setae; 0) two; 1 ) none. 
General ground color of the flies: 0) not yellow; 
1 ) yellow. 



available (Huckett and Vockeroth 1987, 
Carvalho 1989d). 

Table 1 includes all useful characters and 
character states distribution of Soiizalopes- 
myia species and their polarities used in the 
present paper. The program Hennig86, ver- 
sion 1. 5 (Fams 1988) was used for the phy- 
logenetic analysis, applying the implicit 
enumeration (ie*) option. Consistency (CI) 



VOLUME 101. NUMBER 1 



125 



and retention (RI) indices were calculated 
excluding uninformative characters (auta- 
pomorphies and synapomorphies of the ge- 
nus). 

Taxonomy 
Souzalopesmyia Albuquerque 1951 

Peruvia Malloch 1929:104 (preocc. Scud- 
der 1890). Type-species, Mydaea singu- 
laris Stein, 1911 (orig. desig.). 

Souzalopesmyia Albuquerque 1951:53. 
Type-species, Souzalopesmyia carioca 
Albuquerque, 1951 (orig. desig.). 

Diagnosis. — Souzalopesmyia may be rec- 
ognized by its typical head and antenna 
shape, by the setulose parafacials for half 
their length, and by the absence of ocellar 
setae. Also, they are wholly yellow flies ex- 
cept for the presence of stripes on the ab- 
domen (Albuquerque 1951: Figs. 3, 16). 

Description. — Male head dichoptic (Al- 
buquerque 1951: Fig. 1), narrower than in 
female. Frons with orbital setae reclinate, 
and without crossed setae on frontal vitae. 
Ocellar setae absent. Antenna long, reach- 
ing epistome. Arista plumose, longest hairs 
equal to greatest antennal diameter. Para- 
facials setulose on upper half. Female pro- 
boscis as in Fig 14. Dc 1-2:3. Acr not dis- 
tinct from ground setulae, except in female 
of S. amazonica. 2 postpronotals. la: 1:2. 
Sa: 1:2, second weak, about half length of 
first. 2 pa. 2 subequal npl setae. Pra absent. 
Disc of notopleuron bare. Anepisternum 
with a short seta in upper anterior corner. 
Anepimeron, greater ampulla, vallar ridge, 
and meron bare. Ktps 1 :2 (not 1:1:1 as stat- 
ed by Albuquerque 1951). Metathoracic 
spiracle small, triangular, with yellow se- 
tulae on margin. Prosternum bare. Fore tib- 
ia with 1 PD submedian setae. Fore tar- 
somere 1 with 1 V setae. Mid femur with 
1 AD, 1 D, 1 PD and 1 P preapical setae. 
Mid tibia with 2-4 P median setae: 1 
strong V apical setae. Hind tibia without 
calcar and with 1 AD median setae, 1 D, 
1 AD, 1 AV apical setae. Veins bare, ex- 
cept for costa. Vein M , + 2 curved slightly 



toward vein R 4,3. Lower calypter of 
Phaonia-type. Sternite 1 bare. Male aedea- 
gus as in Figs. 10-12. Ovipositor long, ter- 
gites, sternites and membranes covered 
with microtrichia (Figs. 15, 16). Three 
elongate spermathecae. Egg: Phaonia- 
type. 

Monophyly. — Souzalopesmyia Albu- 
querque is a monophyletic genus based on 
the following synapomorphies: 1, Head lat- 
eral appearance elongate; 2, Ocellar setae 
absent; 3, Ground colour yellow. 

Remarks. — The species of Souzalopes- 
myia are rare and have similar facies. Based 
on current collection records, they are 
found in rainforest habitats. They may be 
nocturnal, as they are rarely collected by 
day. 

Key to Species of Souzalopesmyia 

1 Vti divergent. Crossvein dm-cu oblique, 

weakly curved (Albuquerque 1951: Fig. 

13); female: 2 frontals 

S. carioca Albuquerque 

- Vti parallel. Crossvein dm-cu oblique, al- 
most straight (Albuquerque 1951: Fig. 11); 
female: 3 frontals 2 

2(1) Dc 2: 3; male: postocular row of setulae in- 
complete, not reaching epistome; setulae 
black; female: some acr prst stronger than 

the ground setulae 

S. anuizonica Albuquerque 

- Dc 1: 3; male: postocular row of setulae 
complete; setulae black and yellow, the lat- 
ter beginning after basal half of eye; female: 
acr prst undifferentiated from the ground se- 
tulae 3 

3 1 proepisternal seta weak, about % length of 
the upper anepisternal setae; male: fifth ster- 
nite without sharp depression on posterior 
side (Fig. 1); cereal plate heart-shaped (Fig. 

4) S. paraensis Carvalho. new species 

- 1 proepisternal seta strong, similar to the 
upper anepisternal setae; male: fifth sternite 
with sharp depression on posterior side 
(Figs. 2, 3); cereal plate rounded (Figs. 5, 

6) 4 

4 (3) Species ranging from 8.0 to 9.0 mm in 

length; posterior ktps strong, about 2 times 
the length of the anterior one; male: frontal 
setae both cruciate; fifth sternite as in Fig. 

2; cereal plate as in Fig. 5 

5. singiilaris (Stein) 

Species ranging from 6.5 to 8.0 mm in 
length; posterior ktps very strong, about 3 



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times the length of the anterior one; male: 
lower frontal setae cruciate, upper reclinate; 
fifth sternite as in Fig. 3; cereal plate as in 
Fig. 6 S. siilina Carvalho. new species 

Souzalopesmyia amazonica 
Albuquerque 1951 

Souzalopesmyia amazonica Albuquerque 
1951:56; Pont 1972:23 (Neotropical cat- 
alog); Carvalho et al. 1993: 84 (Neotrop- 
ical catalog). 

Diagnosis. — This species is very similar 
to 5. singularis but, it can be easily distin- 
guished from the other Souzalopesmyia 
species by dc 2:3 setae. 

Description. — Male: Head: Frons broad, 
at narrowest point 0.21 of head width. Eye 
with only normal pubescence. Fronto-orbit- 
al plate, parafacial, face and gena silvery 
white. Fronto-orbital plate broad, broaden- 
ing gradually from vertex to lunula; at ver- 
tex plate almost equal to diameter of ante- 
rior ocellus, at lunula equal to three times 
anterior ocellus. Frontal vitta broad, parallel 
to vertex. 2 pairs of strong frontal setae on 
lower % of frons, former cruciate and latter 
reclinate; 1 strong orbital, reclinate and di- 
vergent. Ocellar triangle black, reaching to 
insertion of orbital setae, with some setulae 
behind posterior ocellar setae. Vti strong 
and parallel. Postocular row incomplete, 
reaching as single row just below mid level 
of eye and composed of black setulae. Gena 
below lowest eye margin equal to twice di- 
ameter of the anterior ocellus. Palpi slender, 
yellow. 

Thorax: Ground color yellow, scutum 
dusted with whitish-grey. Dc 2:3; 5-6 rows 
of prst acr setulae; 11-12 rows of post acr 
setulae. 1 proepistemal weak, about % 
length of upper anepisternal setae. Posterior 
ktps strong, about 2.5 times length of an- 
terior one. Scutellum with 1 strong pair of 
apical and 1 subbasal setae; 1 preapical 
weak but stronger than ground setulae; disc 
with setulae descending below strong setae; 
bare ventrally. 

Legs: Yellow. Fore femur with complete 
rows of AV and D setae; AD row weak and 



just to apical %. Mid tibia with 3 P setae. 
Hind tibia with 5 AV setae on apical half 
(Albuquerque 1951: 57). 

Wings: Clear, veins yellow. Membrane 
entirely covered with microtrichia. Cross- 
vein r-m placed before point where vein Rl 
enters costa. Crossvein dm-cu oblique, al- 
most straight. Calypters and haltere yellow. 

Abdomen: Ground-color yellow; in pos- 
terior view; with a narrow black stripe on 
tergite 1 which is enlarged on tergite 3, 4 
and 5 (Albuquerque 1951: Fig. 16). Tergite 
4 and 5 with 2 strong apical setae. 

Terminalia: See Albuquerque 1951: Fig. 
12. 

Measurements: Length of body, 7 mm (n 
= 1). Length of wings, 6 mm (n = 1). 

Female: Differs from male as follows: 
Head: Frons at narrowest point 0.27 of 
head width. Frontal vitta broad, divergent 
to vertex. 3 pairs of strong frontals, former 
2 cruciate. Ocellar triangle black, not reach- 
ing to insertion of 3rd pair of frontals. Some 
acr prst stronger than ground setulae. Cross- 
vein dm-cu oblique, more than in male. 

Terminalia: See Albuquerque 1951: Fig. 
17, 18. 

Measurements: Length of body, 8 mm 
(n= 1). Length of wings, 8 mm (n = 1). 

Remarks. — Adults have been collected in 
the afternoon on flowers and at night. 

Type material examined. — Holotype S in 
MNRJ labelled as follow: "35/Manaus — no 
centro [in the center of the city]/ a noite [at 
night]/15.vi.933 [15 June 1933]/Ant. [An- 
tonio] Paes Filho" [hand label]; Souzalo- 
pesmyia/amazonica sp.n,/30.8.50 [30 Au- 
gust 1950 [examined Albuquerque' date] / 
D. Albuquerque det." [Albuquerque hand 
label]; "Holotipo [holotype]" [red label]. 
Specimen in good condition (Lopes et al. 
1997). Right fore tarsi, median and hind 
legs missing. Segments of the abdomen 
mounted on slide in Canada balsam. 

Other examined material. — Total: 1. 
BRAZIL. Amazonas: Manaus, Ant. Paes 
Filho, 26.V.1933 (1 9 allotype MNRJ). 



VOLUME 101, NUMBER 1 



127 



Souzalopesmyia carioca 
Albuquerque 1951 

Souzalopesmyia carioca Albuquerque 
1951:53; Pont 1972: 23 (Neotropical cat- 
alog); Carvalho et al. 1993: 85 (Neotrop- 
ical catalog). 

Diagnosis. — S. carioca is one of the larg- 
est Souzalopesmyia species and can be dis- 
tinguished from the other species by the di- 
vergent vti. The female has only 2 frontal 
setae. 

Description. — Male: Head: Frons broad, 
at narrowest point 0.24 of head width. Eye 
with only normal pubescence. Fronto-orbit- 
al plate, parafacial, face and gena silvery 
white. Fronto-orbital plate broad, broaden- 
ing gradually from vertex to lunula, at ver- 
tex plate almost equal to diameter of ante- 
rior ocellus, at lunula 3 times diameter of 
anterior ocellus. Frontal vitta broad, parallel 
to vertex. 2 pairs of strong cruciate frontals 
on lower % of frons; 1 strong orbital recli- 
nate and divergent. Ocellar triangle reach- 
ing to insertion of orbital setae. Vti strong, 
divergent. Postoculars in a complete single 
row, reaching to epistome and composed of 
black setulae. Gena below lowest eye mar- 
gin equal to twice diameter of anterior ocel- 
lus. Palpi slender, yellow. 

Thorax: Ground-color yellow with scu- 
tum dusted with whitish grey. Dc 1:3; 6-7 
rows of prst acr setulae; 1 1-12 rows of post 
acr setulae. 1 proepistemal seta very weak, 
about half length of upper anepisternal se- 
tae. Posterior ktps strong, about twice 
length of anterior one. Scutellum with 1 
strong pair of apical and 1 subbasal setae; 
1 preapical weak but stronger than ground 
setulae; disc with setulae descending below 
strong setae; bare ventrally. 

Legs: Yellow. Fore femur with a com- 
plete row of AV, D and AD setae, latter 
weakest. Mid femur with 2-3 PV setae in 
basal half, shorter than femoral depth. Mid 
tibia with 2-3 P setae. Hind femur with 1 
strong preapical AV seta, longer than fem- 
oral depth; AD row complete; 1 PD, 1 D, 



1 AD preapical setae weaker than AV seta. 
Hind tibia with 4 AV setae on apical half. 

Wings: Clear, veins yellow. Membrane 
entirely covered with microtrichia. Cross- 
vein r-m placed before point where vein Rl 
enters costa. Crossvein dm-cu oblique, 
weakly curved in median part. Calypters 
and haltere yellow. 

Abdomen: Ground-color yellow; tergite 3 
and 4 each with blackish markings increas- 
ing in size to posterior margin in posterior 
view; tergite 5 blackish, except hind margin 
(Albuquerque 1951: Fig. 3). A median 
black stripe on tergites 3, 4 and 5. Tergite 
4 and 5 with 2 strong apical setae. 

Terminalia: See Albuquerque 1951: Fig. 
4-10. 

Measurements: Length of body, 8 mm (n 
= 1). Length of wings, 8 mm (n = 1). 

Female: Differs from male as follows: 

Head: Frons at narrowest point 0.27 
times maximum head width. Frontal vitta 
broad, divergent to vertex. 2 pairs of strong 
frontals, cruciate. Ocellar triangle black, not 
reaching insertion of 2nd pair of frontals. 

Terminalia: See Albuquerque 1951: Fig. 
14, 15. 

Measurements: Length of body, 8.5 mm 
(n=l). Length of wings, 8.5 (n = 1). 

Type material examined. — Holotype S in 
MNRJ labelled as follow: "Grajahu/Rio de 
Janeiro/Lopes [Hugo de Souza Lopes]- 
6.1.40 [6 January 1940]"; Souzalopesmyia/ 
carioca sp.n./30.8.50 [30 August 1950] [ex- 
amined Albuquerque date] /D. Albuquerque 
det." [Albuquerque hand label]; "Holotipo 
[holotype]" [red label]. Specimen in good 
condition (Lopes et al. 1997). Right wing 
on a slide mounted attached of pin holo- 
type. Segments of abdomen on a slide 
mounted. 

Other examined material. — Total: 1. 
BRAZIL. Rio de Janeiro: Rio de Janeiro, 
H.S. Lopes, 6.VL1940 (1 9 allotype 
MNRJ). 

Souzalopesmyia paraensis 
Carvalho, new species 

(Figs. 1, 4, 7, 10, 13, 14, 15) 
Diagnosis. — Souzalopesmyia paraensis 
can be distinguished from the other Souz- 



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Figs. 1-6. 1-3, Male fifth sternite, dorsal view. 1. Souzalopesmyia paraensis. 2, 5. singularis. 3, S. sulina. 
4-6, Cereal plate, dorsal view. 4, S. paraensis. 5, S. singularis. 6, S. sulina. 



alopesmyia species by male cereal plate 
rounded in outline and fifth sternite without 
posterior depression. 

Description. — Male: Head: Frons broad, 
at narrowest point 0.21 of head width. Eye 
with only normal pubescence. Fronto-orbit- 
al plate, parafacial, face and gena silvery 
white. Fronto-orbital plate broad, broaden- 
ing gradually from vertex to lunula, at ver- 
tex plate less than diameter of anterior ocel- 
lus, at lunula 3 times diameter of anterior 
ocellus. Frontal vitta broad, parallel to ver- 
tex. 2 pairs of strong cruciate frontal setae 
on lower % of frons; 1 strong orbital recli- 
nate. Ocellar triangle reaching insertion of 
orbital setae. Vti strong, parallel. Postocu- 
lars in a complete single row, reaching ep- 
istome and composed of black and yellow 
setulae; latter beginning after basal half of 



eye and reaching epistome, but 1-2 with 
black setulae. Gena below lowest eye mar- 
gin twice diameter of anterior ocellus. Palpi 
slender, yellow. 

Thorax: Ground-color yellow, dusted 
with whitish grey. Dc 1:3; 7-8 rows of prst 
acr setulae; 10—1 1 rows of post acr setulae. 
1 proepisternal seta weak, about % length 
of the upper anepisternal setae. Posterior 
ktps strong, twice length of anterior one. 
Scutellum with 1 strong pair of apical and 
1 subbasal setae, 1 preapical weak but 
stronger than ground setulae; disc with se- 
tulae descending below strong setae; bare 
ventrally. 

Legs: Yellow. Fore femur with a com- 
plete row of AV and D setae; AD row 
weak. Mid femur with 2 PV setae in basal 
half, not equal to femoral depth. Mid tibia 



VOLUME 101. NUMBER 1 



129 




Figs. 7-12. 7-9. Epandrium. cereal plate and surstylus. lateral view. 7, Souzalopesinyia paraensis. 8. 5. 
singiilaris. 9, S. siilina. 10-12. Phallus and associated structures, lateral view. 10. S. paraensis. 11.5. singiilahs. 
12, S. siilina. 



with 3 P setae. Hind femur with 1 strong 
preapical AV seta, longer than femoral 
depth; AD row complete; 1 PD, 1 D, 1 AD 
preapical seta weaker than AV seta. Hind 
tibia with 4 AV setae on apical half. 

Wings: Clear, veins brownish. Membrane 
entirely covered with microtrichia. Cross- 
vein r-m placed before point where vein Rl 
enters costa. Crossvein dm-cu oblique, al- 
most straight. Calypters and haltere yellow. 

Abdomen: Ground-color yellow; in pos- 
terior view with a narrow, slight, black 
stripe on tergite 1 which is blackish and en- 
larged on tergites 3, 4 and 5. Tergite 4 and 
5 with 2 strong apical setae. 

Terminalia: See Figs. 1, 4, 7, 10, 13. 



Measurements: Length of body, 8.5 mm 
(n = 1). Length of wings, 6.6 (n = 1). 

Female: Differs from male as follows: 
Head: Frons at narrowest point 0.31 of 
head width. Frontal vitta broad, divergent 
to vertex. 3 pairs of strong frontals, first one 
cruciate, second one reclinate and conver- 
gent and third one reclinate and divergent. 
Ocellar triangle black, not reaching inser- 
tion of 3rd pair of frontals. Postoculars in a 
complete single row, with black setulae. 

Measurements: Length of body, 8.5-9.0 
mm (n = 2). Length of wings, 7.0-7.7 mm 
(n = 2). 

Type material examined. — Holotype 6 in 
MPEG, labelled as follow: "Belem Mo- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



cambo/ Ol-IV-1977"; "Brasil Para/A. Y. 
Harada." 

Other examined material. — Total: 2 par- 
atypes: BRASIL. Para: Belem. A. Y. Ha- 
rada, 1. IV. 1977 (1 9 DZUP); ibidem, same 
collector, 6.IV.1977 (1 9 MPEG). Speci- 
men in reasonable condition. Left fore and 
right mid legs missing; left mid tarsi and 
hind right leg glued on to a card attached 
to the pin. 

Souzalopesmyia singidaris (Stein 1911) 
(Figs. 2, 5, 8, 11, 16) 

Mydaea singidaris Stein 1911:91; Stein 
1919: 124 (world catalog); Seguy 1937: 
282 (world catalog). 

Peruvia singularis; Malloch 1929: 105 
(type species of Peruvia)\ Hennig 1965: 
Fig. 31 (tip of female ovipositor). 

Souzalopesmyia singularis; Pont 1972: 24 
(Neotropical catalog); Carvalho et al. 
1993: 85 (Neotropical catalog). 

Diagnosis. — Souzalopesmyia singularis 
can be distinguished from the other Sou- 
zalopesmyia species by posterior depression 
on male fifth sternite and 2 cruciate frontal 
setae. 

Description. — Male: Head: Frons broad, 
at narrowest point 0.23 of head width. Eye 
with only normal pubescence. Fronto-orbit- 
al plate, parafacial, face and gena silvery 
white. Fronto-orbital plate broad, broaden- 
ing gradually from vertex to lunula, at ver- 
tex plate less than diameter of anterior ocel- 
lus, at lunula 2.5 times diameter of anterior 
ocellus. Frontal vitta broad, parallel to ver- 
tex. 2 pairs of strong cruciate frontals on 
lower % of frons; 1 strong orbital reclinate. 
Ocellar triangle reaching insertion of orbital 
setae. Vti strong parallel. Postoculars in a 
complete single row, reaching epistome and 
composed of black and yellow setulae; lat- 
ter beginning after basal half of eye and 
reaching epistome, but 1-2 with black se- 
tulae. Gena below lowest eve margin equal 
to 2.5 times diameter of the anterior ocellus. 
Palpi slender, yellow. 

Thorax: Ground-color yellow with scu- 



tum dusted with whitish-grey. Dc 1:3; 5-6 
rows of prst acr setulae; 9-10 rows of post 
acr setulae. 1 proepisternal seta similar to 
upper anepisternal setae. Posterior ktps 
strong, twice length of anterior ones. Scu- 
tellum with 1 strong pair of apical and 1 
subbasal setae; 1 preapical weak, but stron- 
ger than ground setulae; disc with setulae 
descending below strong setae; bare ven- 
trally. 

Legs: Yellow. Fore femur with complete 
row of AV and D setae; AD row weak. Mid 
femur with 2 PV setae in basal half, less 
than femoral depth. Mid tibia with 3 P se- 
tae. Hind femur with 1 strong preapical AV 
seta, longer than femoral depth; AD row 
complete; 1 PD, 1 D, 1 AD preapical setae 
weaker than AV seta. Hind tibia with 5 AV 
setae on apical half. 

Wings: Clear, veins brownish. Membrane 
entirely covered with microtrichia. Cross- 
vein r-m placed just before point where vein 
Rl enters costa. Crossvein dm-cu oblique, 
almost straight. Calypters and haltere yel- 
low. 

Abdomen: Ground-color yellow; in pos- 
terior view with slight narrow black stripe 
on tergite 1 which is enlarged on tergite 3, 
4 and 5. Tergite 4 and 5 with 2 strong apical 
setae. 

Terminalia: See Figs. 2, 5, 8, 11. 

Measurements: Length of body, 8 mm (n 
= 1). Length of wings, 7.5 mm (n = 1). 

Female: Differs from the male as fol- 
lows: Head: Frons at narrowest point 0.32 
times maximum head width. Frontal vitta 
broad, divergent to vertex. 3 pairs of strong 
frontals, first one cruciate, second one rec- 
linate and convergent and third one recli- 
nate and divergent. Ocellar triangle not 
reaching insertion of 3rd pair of frontals. 
Postoculars in a complete single row of 
black setulae. 

Measurements: Length of body, 8.5-9.0 
(n = 3). Length of wings. 8.0-8. 5 mm (n 
= 3). Type material examined. Holotype 
male in SMT labelled as follow: "Bolivia- 
Mapiri/ 14. III. 03 [hand written]/Sarampioni 
700 m [green label]": "Coll W. Schnuse/ 



VOLUME 101. NUMBER 



131 




r 








15 




Figs. 13-16. 13-15, Souzalopesmyia paraensis. 13. Hypandrium, dorsal view. 14, Proboscis, lateral view. 
15. Ovipositor, ventral view. 16. Ovipositor of 5. singiilohs. ventral view. 



1911-3"; "Mydaea/singularis/ sp.n. [light 
green hand written label]: "Typus" [red la- 
bel]; "Staatl. Museum fur/ Tierlunde Dres- 
den." Specimen in good condition lacking 
the left hind leg and hind right tarsi. Ab- 
domen in microvial with glycerine. 

Other examined material. — Total: 3. BO- 
LIVIA: Mapiri, Sarapioni, 700-800 m, W. 



Schnuse, III- 1903 (2 9 SMT); S. Carlos, W. 
Schnuse, 1-1901 (1 9 SMT). 

Souzalopesmyia sulina 
Carvalho, new species 

(Figs. 3, 6, 9, 12) 

Diagnosis. — Souzalopesmyia sulina is 
one of the smallest Souzalopesmyia species. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



It can be distinguished from the other Souz- 
alopesmyia species by deeply posterior de- 
pression on male fifth sternite and 2 frontal 
setae, latter reclinate. 

Description. — Male: Head: Frons broad, 
at narrowest point 0.25 of head width. Eye 
with only normal pubescence. Fronto-orbit- 
al plate, parafacial, face and gena silvery 
white. Fronto-orbital plate broad, broaden- 
ing gradually from vertex to lunula, at ver- 
tex plate about equal to diameter of anterior 
ocellus, at lunula 2.5 times diameter of an- 
terior ocellus. Frontal vitta broad, parallel 
to vertex. 2 pairs of strong frontal setae on 
lower % of frons; former cruciate and latter 
reclinate; 1 strong orbital reclinate and di- 
vergent. Ocellar triangle black, reaching in- 
sertion of orbital setae. Vti strong and par- 
allel. Postoculars in a complete single row, 
reaching epistome and composed of black 
and yellow setulae; latter beginning after 
basal half of eye and reaching epistome, but 
1-2 with black setulae. Gena below lowest 
eye margin twice diameter of anterior ocel- 
lus. Palpi slender, yellow. 

Thorax: Ground-color yellow with scu- 
tum dusted with whitish-grey, more evident 
in pre sutural area. Dc: 1:3; 5-6 rows of 
prst acr setulae: 11-12 rows of post acr se- 
tulae. 1 proepistemal seta strong, similar to 
upper anepisternal setae. Posterior ktps very 
strong, about 3 times length of anterior one. 
Scutellum with 1 strong pair of apical and 
1 subbasal setae; 1 preapical weak but 
stronger than ground setulae; disc with se- 
tulae descending below strong setae; bare 
ventrally. 

Legs: Yellow. Fore femur with complete 
rows of AV and D setae; AD row weak. 
Mid femur with 2-3 PV setae in basal half, 
less than femoral depth. Mid tibia with 3- 
4 P setae. Hind femur with 1 strong preap- 
ical AV seta, longer than femoral depth; AD 
row weak and complete; 1 PD, 1 D, 1 AD 
preapical seta weaker than AV seta. Hind 
tibia with 4-5 AV setae on apical half. 

Wings: Clear, veins brownish. Membrane 
entirely covered with microtrichia, cross- 
vein r-m placed just before point where vein 



Rl enters costa. Cross vein dm-cu oblique, 
almost straight. Calypters and haltere yel- 
low. 

Abdomen: Ground-color yellow; in pos- 
terior view with a narrow black stripe on 
tergite 1 which is enlarged on tergite 3, 4 
and 5. Tergite 4 and 5 with 2 strong apical 
setae. 

Terminalia: See Figs. 3, 6, 9, 12. 

Measurements: Length of body, 6.5-8.0 
mm (n = 5). Length of wings, 6.5-7.7 mm 
(n = 5). 

Female: Unknown. 

Remarks. — Adults have been collected 
with Malaise traps. 

Type material examined. — Holotype 6 in 
MHPA labelled as follow; "Depto Caninde- 
yu/Reserva Natural del Bosque/Mbaracayu: 
Jejui-mi/Malaise 3, bosque bajo inundado/ 
Coh-. A.C.F Costa/ 10-18.VIV1996 [18 July 
1996]"; "Holotipo [holotype]" [red label]. 

Other material examined. — Total: four 
paratypes. Same label of the holotype: 2 S, 
10-18 July 1996 [DZUP MHPA], 2 c?, 18- 
28 July 1996 [DZUP, MHPA]. 

Discussion 
Phylogenetic Analysis 

Phylogenetic studies on Muscidae are 
still scarce (Carvalho 1989d). The family 
contains about 200 genera with well over 
4,000 species worldwide (Pont 1989, Car- 
valho et al. 1993). The family is undoubt- 
edly monophyletic (Hennig 1965, Mc- 
Alpine 1989, Michelsen 1991), but histori- 
cally the Muscidae has included groups that 
are doubtfully monophyletic (Carvalho 
1989d). 

Several genera of Phaoniinae in the Neo- 
tropics (Pont 1972), a paraphyletic subfam- 
ily pointed by Hennig (1965) and recently 
by Carvalho (1989b), were transferred to 
other subfamilies (Carvalho 1985, 1989a, 
1989b, 1989c, Couri and Lopes 1986, Car- 
valho & Pont 1998). Currently the subfam- 
ily in the region has only four genera (Car- 
valho et al. 1993): Dolichophaonia Carva- 



VOLUME 101. NUMBER 1 133 

S amazonica S. paraensis, sp n S singularis S sulina, sp. n. S. carioca 



10 



11 



6,7 



8, 10 



17 



Fig. 17. Cladogram of species of Souzalopesmyici. Synapomorphies = solid black rectangles: reversal 
dotted rectangles. Character numbers correspond to those stated in Table 1. 



Iho, 1993; Helina, Phaonia and Souzalo- 
pesmyici. 

Souzalopesmyia is a small and isolated 
clade, apparently representing an ancient 
lineage of Phaoniinae. The species of genus 
have head elongate which is commonly cor- 
related with broad male frons (Vockeroth 
1972). But these lengthening is considered 
independent from that in the some genera 
in the subfamilies Atherigoninae, Cyrtoneu- 
rininae, Azeliinae-Reinwardtiini or Myda- 
einae. 

The Hennig86 phylogenetic analysis of 
Souzalopestnyia was based on seven char- 
acters (with an asterisk in Table 1) and re- 
sulted in a single tree shown in Fig. 17 
(length = 9 steps, consistency index = 
0.77, retention index = 0.75). 



Biogeography 

Hennig (1965) argued that the first in- 
vasion (Edentata level) of the Neotropical 
Muscidae fauna may have come from the 
Northern Hemisphere during the Upper 
Cretaceous or Early Tertiary period. This 
time frame was also suggested by Michel- 
sen (1991) for the invasion of the basal 
clade of the Anthomyiidae, Coenosopsia + 
Phaonantho, into South America. 

There are few papers on biogeography of 
Muscidae, none of them of Neotropical Re- 
gion, except Hennig (1965). In Holarctic 
Region, all taxa of the Eudasyphora s. str. 
Townsend (Muscinae) are faunal elements 
of known dispersal centers (Cuny 1980). 
The speciation of these flies, were correlat- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



n 


4- 

\J ^ Caribbean Plate 




^w 


^^IJ 






k^ 


Separation of the 
Northwestern and the 
^ Southeastern Components 


Vicanance of 
Northwestern 
Amazonia 


^▼^N(^^ v>_^ 


fe- 




if ^ 


^f rK. Mantiqueira 




Andes y M ^ 


^ 18 



Fig. 18. A summary of the main vicariance baniers in the history ot the Neotropical region. 1 = Cairibbean 
Plate detachment from the mainland. 2 = Separation between the Northwestern and the Southeastern main 
components, a line along the Rivers Amazonas/Madeira/Mamore in the Amazonian Basin. 3 = Epicontinental 
sea formation in the Maracaibo area. 4 = A large division in northwestern Amazonia (not related to date to any 
geological event). 5 = Middle to Late Cretaceous water connection between the Parnafba Basin and the Parana 
Basin (redrawn from Amorim and Pires 1996). 



ed with the history of the forest vegetation 
during the Pleistocene (Cuny 1980). 

Amorim and Pires (1996), corroborated 
independently by Grazia (1997), indicated 
that the first division in the continental re- 
gion of the Neotropics was in the Late Cre- 
taceous, showing a northwestern track 
against a southeastern track (Fig. 18). Alike 
pattern is showed by Camargo (1996) for 
some bees (Meliponini, Apinae, Apidae) in 
Neotropical Region. However, this latter 



biogeographical reconstruction was postu- 
lated by modification occurred by the 
changing forests in the Pleistocene. 

Souzalopesmyia, based on the position of 
S. amazonica as the basal clade of genus, 
may have had its ancestor back in the Late 
Cretaceous (Fig. 19). This age for the genus 
is not unrealistic although no fossil record 
is known for family older than the Eocene 
(Evenhuis 1994). Pont and Car valho (1997) 
studied three fossil species of Muscidae 



VOLUME 101, NUMBER 1 



135 




Fig. 19. Geographical distribution of species of Soiizalopesmyici, with cladogram superimposed, a, S. ama- 
zonica. b. 5. paraensis. c, S. singulans. d. S. sitlina. e. S. carioca. 



136 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



from Dominican amber dated from Mio- 
cene, 15-20 mya ago, two of which are 
Phaonia species. A cladogram by S. M. P. 
Coelho (unpublished Ph.D. thesis) suggests 
that the Phaonia fossil species have a more 
recent origin than the ancestor of Souzalo- 
pesmyia. The origin of the genus is there- 
fore probably older than 15-20 mya, sug- 
gesting that Soiizalopesmyia is one of the 
most basal genera of Phaoniinae in the Neo- 
tropics. 

The origin of the Neotropical Phaoniinae 
fauna cannot be completely understood on 
the basis of the present paper. The ancestor 
of Soiizalopesmyia may have reached South 
America by dispersal from North America 
(Hennig 1965), Africa or have evolved in 
the Neotropics. The discovery of the sister 
group of the genus is required. 

The five species of Soiizalopesmyia are 
morphologically similar and exhibit allo- 
patric distribution suggesting that the ter- 
minal branches of this clade could be re- 
sulted from relatively recent speciation. 

Nevertheless, the allopatric pattern of 
speciation of the genus cannot be fully ex- 
plained with the available geological and 
biogeographical information. Most of the 
species are known from only a few speci- 
mens, suggesting that intensive effort may 
be necessary before reliable statements can 
be made about the distribution patterns of 
the species. Species of Soiizalopesmyia are 
not expect to occur in western side of An- 
des as it requires tropical rainforest (Figs. 
18, 19). 

The occurrence of S. paraensis in Belem, 
along south side of the Amazon River, part 
of the northwestern track, could be the re- 
sult of a single dispersal event (Figs. 18, 
19) to colonize that region, in a more recent 
time, which belongs to the northwestern 
track. 

Acknowledgments 

I express my sincere gratitude to the fol- 
lowing persons who supplied specimens for 
this study: Uwe Kallweit (SMT), D.M. 
Pamplona (MNRJ), T Pimentel (MPEG) 



and John Kochalka (MNPA). I also thank 
A. C. F. da Costa, The Fundacion Moises 
Bertoni and the Darwin Initiative for logis- 
tic support for the project Biodiversity In- 
ventory of the Mbaracayu Forest Nature 
Reserve, Paraguay. I thank the following 
for critically reviewing the manuscript: L. 
Marinoni, Universidade Federal do Parana, 
Curitiba, Brazil; D. S. Amorim, Universi- 
dade Sao Paulo, Ribeirao Preto, Brazil; M. 
S. Couri and D. M. Pamplona. Museu Na- 
cional do Rio de Janeiro, Rio de Janeiro, 
Brazil; G. S. Carvalho, Pontificia Univer- 
sidade Catolica, Porto Alegre, Brazil; A. Y. 
Harada, Museu Paraense Emflio Goeldi, 
Belem, Brazil; A. C. Pont, University Mu- 
seum of Natural History, Oxford, U. K; R. 
Holzenthal, University of Minnesota, 
U.S.A. I am deeply grateful to R. Cuny, 
Lakeland College, Alberta, Canada, for 
comments and additions, which have mark- 
edly improved the paper. This study was 
partially supported from CNPq, Conselho 
Nacional de Desenvolvimento Cientifico e 
Tecnologico (300.043/86-4). This is paper 
number 1035 of the Departamento de Zoo- 
logia, UFPR. 

Literature Cited 

Albuquerque, D. de O. 1951. Sobre um genero e duas 
especies novas de "Phaoniinae" neotropical (Di- 
ptera, Muscidae). Revista Brasileira de Biologia 
11: 53-57. 

Amorim, D. de S. and M. R. S. Pires. 1996. Neotrop- 
ical biogeography and a method for maximum 
biodiversity estimation, pp. 183-219. In Biodiver- 
sity in Brasil, a First Approach. CNPq, Sao Paulo. 
326 pp. 

Camargo, J. M. F. 1996. Meliponini neotropicais (Ap- 
inae, Apidae, Hymenoptera): Biogeografia Histo- 
rica. pp. 107-121. In Anais do II Encontro sobre 
abelhas. Ribeirao Preto, 351 pp. 

Carvalho, C. J. B. de. 1985. A new systematic position 
for Scenetes Malloch, 1936 with a description of 
the genitalia of 5. cardini Malloch (Diptera, Mus- 
cidae, Mydaeinae). Revista Brasileira de Ento- 
mologia 29: 575-577. 

. 1989a. Revisao das especies e posigao siste- 

matica de Palpibracns Rondani (Diptera, Musci- 
dae). Revista Brasileira de Zoologia 6: 325-376. 

. 1989b. Revisao dos generos sul-americanos: 

Brachygasterina Macquart e Conentosia Malloch 



VOLUME 101. NUMBER 1 



137 



(Diptera, Muscidae). Revista Brasileira de Zoo- 

logia 6: 473-484. 
. 1989c. Revisao de Psilochaeta Stein e des- 

cri^ao de Dalcyella gen.n. do Chile (Diptera, 

Muscidae). Revista Brasileira de Zoologia 6: 485- 

506. 
. 1989d. Classifica9ao de Muscidae (Diptera): 



uma proposta atraves da analise cladi'stica. Revista 
Brasileira de Zoologia 6: 627-648. 

Carvalho, C. J. B. de, M. S. Couri, A. C. Pont, D. 
Pamplona and S. M. Lopes. 1993. Part IL Mus- 
cidae, 201 pp. In Carvalho, C. J. B. de, ed., A 
Catalogue of the Fanniidae and Muscidae (Di- 
ptera) of the Neotropical Region. Sao Paulo, So- 
ciedade Brasileira de Entomologia. 

Carvalho, C. J. B. de and A. C. Pont. 1998. A revision 
of New World Brontaea Kowarz (Diptera, Mus- 
cidae). Revista Brasileira de Zoologia 14 [1997]: 
723-749. 

Couri, M. S. and S. M. Lopes. 1986. Neotropical gen- 
era of Coenosiinae — Nomenclatural notes and key 
to identification (Diptera-Muscidae). Revista 
Brasileira de Biologia 45[1985]: 589-595. 

Cuny, R. 1980. Revision of the genus Eudasyphora 
Townsend (Diptera: Muscidae), and reflections on 
its evolution. The Canadian Entomologist 112: 
345-373. 

Evenhuis, N. L. 1994. Catalogue of the fossil flies of 
the world (Insecta: Diptera). Backhuys. 600 pp., 
Leiden. 

Farris, J. S. 1988. Hennig86 reference. Version 1.5. 
Published by the author. New York. 

Grazia, J. 1997. Cladistic analysis of the Evoplitus ge- 
nus group of Pentatomini (Heteroptera: Pentatom- 
idae). Journal of Comparative Biology 2: 43-47. 

Hennig, W. 1965. Vorarbeiten zu einem phylogene- 
tischen System der Muscidae (Diptera: Cyclor- 
rhapha). Stuttgarter Beitraege zur Naturkunde 
141: 100 pp. 

Huckett, H. C. and J. R. Vockeroth. 1987. Muscidae, 
pp. 1115-1131. //; McAlpine, J. F, B. V. Peterson. 
G. E. Shewell, H. J. Teskey, J. R. Vockeroth and 
D. M.Wood, eds.. Manual of Nearctic Diptera, 2. 
Research Branch Agriculture. Canada Monograph 
28: 675-1332. 

Lopes, S. M., M. Couri, D. Pamplona, and C. J. B. de 
Carvalho. 1997. Notes on Neotropical types of 
Diptera described by Albuquerque (Anthomyidae 
[sic], Fanniidae, Muscidae, Piophilidae, Psilidae, 
Sapromyzidae, Scatophagidae e Stratiomyidae). 



Publica96es Avulsas do Museu Nacional, Rio de 
Janeiro, n. 69: 1-33. 

Malloch, J. R. 1929. Exotic Muscaridae (Diptera). — 
XXVI. Annals and Magazine of Natural History 
(10) 4: 97-120. 

McAlpine, J.F. 1981. Morphology and terminology, pp. 
9-63. In McAlpine, J. F, B. V. Peterson, G. E. 
Shewell, H. J. Teskey, J. R. Vockeroth and D. 
M.Wood, eds.. Manual of Nearctic Diptera, 1. Re- 
search Branch Agriculture. Canada Monograph 
27: 1-674. 

Michelsen, V. 1991. Revision of the aberrant New 
World genus Coenosopsia (Diptera: Anthomyi- 
idae), with a discussion of anthomyiid relation- 
ships. Systematic Entomology 16: 85-104. 

O'Hara, J. E. 1982. Classification, phylogeny and zoo- 
geography of the north American species of Si- 
phona Meigen (Diptera: Tachinidae). Quaestiones 
Entomology 18: 261-380. 

Pont, A. C. 1972. Family Muscidae. //; A Catalogue 
of the Diptera of the Americas South of the United 
States. 97. Museu de Zoologia. Universidade de 
Sao Paulo. Ill pp. 

. 1980. Family Muscidae, pp. 721-761. In 

Crosskey, R. W, ed.. Catalogue of the Diptera of 
the Afrotropical Region. British Museum (Natural 
History). London. 1437 pp. 

. 1986. Studies on Australian Muscidae (Dip- 
tera) vn. The genus Atherigona Rondani. Austra- 
lian Journal of Zoology, Supp. Sen, 120: 1-90. 

. 1989. Family Muscidae, pp. 675-699. //; Ev- 



enhuis, N., ed.. Catalog of the Diptera of the Aus- 
tralasian and Oceanian Regions. Bishop Museum 
Press, Honolulu & E.J. Brill, Leiden. 1155 pp. 

Pont. A. C. and C. J. B de Carvalho. 1997. Three spe- 
cies of Muscidae (Diptera) from Dominican am- 
ber. Studia Dipterologica 4: 173-181. 

Seguy, E. 1937. Diptera Family Muscidae. //) Wyts- 
man. P., Genera Insectorum. Ease. 205: 604 pp., 
Bruxelles. 

Stein, P. 1911. Die von Schnuse in Siidamerika gefan- 
genen Anthomyiden. Archiv fiir Naturgeschichte 
77 (1): 61-189. 

. 1919. Die Anthomyidengattungen der Welt, 

analytisch bearbeitet, nebst einem kritisch-syste- 
matischen Verzeichnis aller aussereuropaische Ar- 
ten. Archiv fiir Naturgeschichte 83 A 1 [1917]: 
85-178. 

Vockeroth, J. R. 1972. A Review of the World Genera 
of Mydaeinae, with a Revision of the Species of 
New Guinea and Oceania (Diptera: Muscidae). 
Pacific Insects Monograph 29: 134 pp. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999, pp. 138-142 

ASHLOCKOBIUS, A NEW GENUS OF MYODOCHINI FROM VENEZUELA 
(HEMIPTERA: LYGAEOIDEA: RHYPAROCHROMIDAE: MYODOCHINI) 

James A. Slater and Alex Slater 

(JAS) Department of Ecology and Evolutionary Biology, University of Connecticut, 
Storrs, CT 06269, U.S.A.; (AS) 901 Alabama Street, Lawrence, KS 66044, U.S.A. 



Abstract. — Ashlockobius cursorius, new genus and species of myodochine lygaeoid 
is described from Venezuela. The genus is related cladistically to Orthaea Dallas, Catenes 
Distant, Heraeiis Stal and Myodocha Latreille. It is believed to be an ant mimic. Figures 
are given of the details of the inflated phallus and of the claspers. A color figure of the 
habitus is included. 

Key Words: Hemiptera, Lygaeoidea, Rhyparochromidae, Myodochini. Venezuela, Ash- 
lockobius cursorius, mimicry 



The Myodochini constitute one of the 
dominant elements in the lygaeoid fauna of 
the Neotropics. It is abundant and diverse 
not only in the number and variety of taxa 
present but in the abundance of many in- 
dividuals of some species. Ant mimicry oc- 
curs frequently. The new genus described 
below is apparently an ant mimic although 
its biology is unknown. 

Henry (1997) has elevated the former 
subfamily Rhyparochrominae of the family 
Lygaeidae to family status with two sub- 
families, the Plinthisinae and the Rhyparo- 
chrominae with the former tribes of the lat- 
ter retained as tribes. We have adopted Hen- 
ry's conclusions here, but suggest that the 
tribes within his new definition of the sub- 
family may ultimately prove to merit family 
status cladistically. Henry's statement that 
the subfamily Rhyparochrominae as rede- 
fined consists of taxa with an incomplete 
suture between abdominal sterna four and 
five and with carinate pronotal margins is 
an oversimplification. Genera that occur in 
at least five tribes within the Rhyparo- 
chrominae have the abdominal sternal su- 
ture complete. Rounded, ecarinate pronota 



occur in several tribes and is one of the di- 
agnostic features of the Myodochini. 
All measurements are in millimeters. 

Ashlockobius Slater and Slater, 
new genus 

Description. — Body elongate, slender, 
nearly parallel sided. Dorsal surface of 
body subshining. Legs and antennae both 
extremely elongate. Head moderately decli- 
vent anteriorly; eyes set a short distance 
away from anterior pronotal angles. Anten- 
niferous tubercules divergent. Head below 
broadly transversely striate from level of 
distal ends of antenniferous tubercles to 
level of posterior margins of eyes. Bucculae 
V-shaped. Vertex convex between eyes with 
patches of pruinosity present. 

Anterior pronotal lobe extremely elon- 
gate and elliptically convex, with a pruinose 
median stripe on anterior pronotal lobe and 
large irregular pruinose patches laterally on 
posterior half of anterior lobe. Posterior 
pronotal lobe dull with a patch of white 
pruinosity near transverse impression on ei- 
ther side of midline. Pronotum with only 
scattered, relatively inconspicuous, punc- 



VOLUME 101. NUMBER 1 



139 



tures present; anterior collar complete, 
coarsely punctate, delimited posteriorly by 
a sharp deeply impressed line, not produced 
posteriorly at meson; transverse pronotal 
impression deep and complete; posterior 
pronotal lobe with prominent punctures. 

Scutellum with a conspicuous Y-shaped, 
elevated carina; pruinose, bicolored with a 
large triangular basal grayish-white patch 
and a spot on each side near divergence of 
elevated Y-carina, remainder of scutellum 
reddish brown. Clavus, corium and mem- 
brane contrastingly strongly shining former 
with 3 complete rows of punctures and a 
partial fourth row on distal half between in- 
ner and median rows. Corium moderately 
expanded posteriorly; outer margin irregu- 
lar and beaded, lacking a stridulitrum. Api- 
cal corial margin adjacent to membrane 
lacking a series of punctures. Mesepimeron 
not emergent. Metathoracic scent gland au- 
ricle short, straight, tapering distally; evap- 
orative area large, covering inner two thirds 
to three fourths of metapleuron, and ex- 
tending narrowly along posterior margin of 
mesopleuron. 

Fore coxa with a prominent spine and a 
smaller secondary spine present. Fore fe- 
mur elongate and slender, almost entire 
ventral surface of each femur heavily spi- 
nous, with rows of large spines along inner 
and outer edges and numerous small spines 
between. Each fore tibia with a widely 
spaced series of 4 short sharp spines on in- 
ner face. Shaft of tibia not strongly curved. 

Lateral and ventral surfaces completely 
dull and chiefly pruinose except for a large 
quadrate shining patch mesally on meso- 
sternum, this latter with a narrow median 
groove. Antenna very elongate, slender, te- 
rete with fourth segment fusiform. Posterior 
margin of abdominal sternum 2 not finely 
scalloped. 

Posterior margin of pygophore broadly 
rounded, without a median impression. 
Clasper (Fig. 3) with a distinct thumb-like 
exterior projection, interior margin with a 
pronounced flange narrowing distally but 
extending almost to apex. Phallus (Figs. 2, 



4) without sclerotized conjunctival or vesi- 
cal spines; conjunctiva short, beaiing low 
lateral subapical lobes; vesica short; heli- 
coid process present; a large lobe bearing 
many projections on each side of ejacula- 
tory reservoir. Ejaculatory reservoir with 
well developed wings (Fig. 2), holding 
sclerites short, narrowing distally. 

Type species. — Ashlockobius cursorius, 
new species. Monotypic. 

Discussion.- — Ashlockobius keys without 
difficulty to couplet 51 in Harrington's 
(1980) key to myodochine genera. It differs 
from Togo Bergroth (from Japan) in having 
a relatively much longer anterior pronotal 
lobe, more than one and one-half times the 
length of the posterior lobe in Ashlocko- 
bius; in Togo the anterior lobe is at most 
only slightly longer than the posterior lobe. 
Ashlockobius also does not have strongly 
curved anterior tibiae. Most specimens of 
Togo are brachypterous, the male fore tibiae 
are prominently curved, and the members 
of the genus are relatively stout and robust. 

Ashlockobius differs from those genera 
reached through couplet 52 of Harrington 
by having the combination of armed male 
fore tibiae, as described above, and an an- 
terior pronotal lobe at least 1.5 times as 
long as the posterior lobe. 

The absence of vesical or conjunctival 
spines and the robustly winged ejaculatory 
reservoir with short holding sclerites places 
Ashlockobius in the group of genera with 
Harrington's Type IV phallus. This place- 
ment is supported by the broadly rounded 
posterior margin of the pygophore (Fig. 5). 
Within this group the presence of pruinose 
areas on the dorsum and the somewhat 
elongate head and rounded vertex place the 
genus with a group on Harrington's clado- 
gram consisting of Orthaea Dallas, Catenes 
Distant, Heraeus Stal, and Myodocha La- 
treille. Analysis of other phallic characters 
must await study of the inflated phallus of 
more species. The illustration of an inflated 
Myodocha phallus provided by Ashlock 
( 1957), which exhibits an elongate conjunc- 
tiva with several lobes and an apparently 



140 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. \. Ashlockohius ciirsorius. Dorsal view. 



VOLUME 101. NUMBER 



141 




Figs. 2-5. Ashlockobius ciirsorius. 2. Ejaculatory reservoir, dorsal view. 3, Claspers. outer and inner views. 
4, Aedeagus, lateral view. 5, Genital capsule, dorsal view. 



asymmetric set of vesical lobes, indicates 
that such studies will prove to be fruitful. 

Etymology. — It gives us great pleasure to 
be able to dedicate this striking new genus 
to the memory of Dr. Peter D. Ashlock for 
his many contributions to the systematics of 
the Lygaeoidea. 



Ashlockobius cursorius Slater and 
Slater, new species 

(Figs. 1-5) 
Description. — Male: Body very elon- 
gate, slender, attenuated, with extremely 
elongate appendages. Color bright reddish 
brown almost throughout, including ap- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



pendages. Antenna with distal ends of seg- 
ments II and III infuscated with chocolate 
brown. Corium marked with chocolate 
brown as follows: a small spot along lateral 
corial margin at level of distal third of api- 
cal corial margin, a small apical spot, an 
elongate dash along radial vein running 
from level of distal fifth of claval commis- 
sure nearly to anterior end of apical corial 
margin, a stripe at inner angle of corium 
that extends from inner margin at apical 
third of claval commissure to adjacent row 
of punctures then narrowly along apical 
corial margin to a level slightly posterior to 
lateral corial spot. Anterior collai" and pos- 
terior pronotal lobe yellow, contrasting with 
red-brown anterior lobe. Corium light yel- 
low brown except as noted above. 

Eyes large, protrudant but not stalked, 
head strongly narrowing behind eyes but 
without a distinct stalked neck. Length head 
1.34, width 1.20, interocular space 0.62. 
Length pronotum 2.60, length anterior 
pronotal lobe (less anterior collar) 1.60, 
width 1.84. Length scutellum 1.24, width 
0.96. Length claval commissure 0.84. Mid- 
line distance apex clavus-apex corium 1 .76. 
Midline distance apex corium-apex abdo- 
men 1.72. Length labial segments I 0.76, II 
0.84, III 0.52, IV 0.44. Labium reaching 
posterior third of prosternum but remote 
from fore coxae. Length antennal segments 
I 1.88, II 3.48, III 2.80, IV 1.72. Total body 
length 9.80. 



Holotype.— (?. VENEZUELA: Aragua, 
16 km S. Telerias, May 10, 1978 (C.W. & 
L.B. O'Brien and Marshall). In American 
Museum of Natural History, New York. 

Etymology. — The name is from the Latin 
word for a runner. 

Discussion. — The male specimen upon 
which the preceding description is based 
was collected in Venezuela twenty years 
ago. It is a striking, elongate, long-legged 
myodochine, which we have held for many 
years in the hope that additional specimens 
would become available. Unfortunately this 
has not happened. 

Ac KNO WLEDGMENTS 

We thank Drs. Charles and Lois O'Brien 
(Florida State University, Tallahasee) for 
the gift of the holotype and Mrs. Mary Jane 
Spring (University of Connecticut) for ex- 
ecution of the dorsal view illustration. 

Literature Cited 

Ashlock. P. D. 1957. An Investigation of the Taxonom- 
ic value of the Phallus in the Lygaeidae (Hemip- 
tera-Heteroptera). Annals of the Entomological 
Society of America 50; 407-426. 

Hanington, B. J. 1980. A generic level revision and 
cladistic analysis of the Myodochini of the World 
(Hemiptera, Lygaeidae, Rhyparochrominae). Bul- 
letin of the American Museum of Natural History 
167: 49-116. 

Henry. T J. 1997. Phylogenetic Analysis of Family 
Groups within the Infraorder Pentatomomorpha 
(Hemiptera: Heteroptera). with Emphasis on the 
Lygaeoidea. Annals of the Entomological Society 
of America 90: 275-301. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999, pp. 143-148 

BIOSYSTEMATIC STUDIES OF CEYLONESE WASPS, XXII: 

BETHSMYRMILLA, A NEW GENUS OF MUTILLID WASPS 

(HYMENOPTERA: MUTILLID AE: MYRMILLINAE) 

Karl V. Krombein and Arkady S. Lelej 

(KVK) Department of Entomology, MRC 165, National Museum of Namral History, 
Smithsonian Institution, Washington, DC 20560-0165, U.S.A.; (ASL) Institute of Biology and 
Pedology, Vladivostok-22, 690022, Russia (e-mail: entomol@online.marine.su) 



Abstract. — Bethsmyrmilla alticola, new genus and new species is described from Sri 
Lanka. A key is given to the Oriental genera of Myrmillinae. The halictine bee Lasio- 
glossiim (Sudila) alphemim (Cameron) is the probable host of B. alticola. 

Key Words: Mutillid wasps, Bethsmyrmilla alticola, new genus and new species, Sri 
Lanka, Lasioglossmn (Sudila) alphenum (Cameron) 



Borge Petersen estimated that the mutil- 
lid fauna of Sri Lanka was 74 species based 
on the literature (in litt. to KVK, 1976). He 
mentioned that he found 45-50 species 
among the specimens that he had borrowed. 
The latter figure, however, does not incor- 
porate a study of all of the rich mutillid 
fauna collected during the later years, 
1975-1981, of the Smithsonian's "Ceylon 
Insect Project" and two visits in 1993 and 
1997 by K. V. Krombein and B. B. Norden. 
We anticipate that a study of these collec- 
tions will result in a much more accurate 
inventory of the mutillid fauna. 

During the latest trip to Sri Lanka, Beth 
Norden found two females of an unusual 
small mutillid wasp in the ground nests of 
the halictine bee Lasioglossum (Sudila) al- 
phemim (Cameron). These wasps belong to 
a new genus and species that we place in 
the Myrmillinae because the side of the tho- 
rax is evenly concave, the mesopleuron has 
a strong supracoxal carina and the pronotal- 
mesopleural suture is lacking except a small 
section above. The subfamily presently 
comprises three genera in the Oriental re- 



gion: Spilomutilla Ashmead, Squamulotilla 
sensu Mickel and Bethsmyrmilla, new ge- 
nus. 

The genus Squamulotilla Bischoff (type 
species Squamulotilla denticollis Bischoff, 
male. North Cameroon, Nigeria) includes 
seven Afrotropical species which are 
known from males only. For many years 
Guido Nonveiller has collected and studied 
the mutillid fauna of Cameroon. We support 
his opinion (1995) that true females of 
Squamulotilla are probably described in the 
Afrotropical genera Clinotilla Arnold 
(males still unknown). Oriental species 
placed in Squamulotilla by Mickel (1933, 
1935) comprise several undescribed genera 
of Myrmillinae which will be treated sep- 
arately. 

Depositories for specimens listed are as 
follows: 

IBPV Institute of Biology and Pedology, 
Vladivostok, Russia. 

USNM National Museum of Natural His- 
tory, Smithsonian Institution, 
Washington, D.C. U.S.A. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Key TO THE Oriental Genera of 
Subfamily Myrmillinae 

1 . Female 2 

- Male (unknown for Bethsniynnilla) 4 

2. Mandible widened apically, without subbasal 
tooth on inner margin (Fig. 8); gena dentate 
beneath; gastral lergum 2 posteriorly with three 
spots (Fig. 3) or a wide band of pale pubes- 
cence shallowly concave medially; gastral ster- 
num 2 with median longitudinal carina ending 
usually in an acute tubercle 

Spiloniiililla Ashniead 

- Mandible not widened apically, with subbasal 
tooth on inner margin (Figs. 6, 7, 9); gena not 
dentate beneath; gastral tergum 2 posteriorly 
with a band of pale pubescence that widens 
medially to an obtuse angle (Figs. 1, 2, 5) or 
with a large median spot (Fig. 4); gastral ster- 
num 2 if carinate not ending in a tubercle ... 3 

3. Mid trochanter with narrow apical process 
(Fig. 12); posterolateral angles of head tuber- 
culate behind eyes (Figs. 14, 15); thorax gently 
sloping posteriorly, upper margin not dentate 
nor serrate (Figs. 15, 16); mandible apically 
with a single tooth (Fig. 6) 

. . Bethsmynnilla Krombein and Leiej, new genus 

- Mid trochanter without process; posterolateral 
angle of head not tuberculate but gena tuber- 
culate below eye in Squaimtlotilla lamellatci 
Mickel and S. arunciiiuicea Pagden; thorax 
abruptly sloping posterorly, upper margin usu- 
ally serrate or dentate, rarely with only an acute 
median denticle, or if not dentate at all (S. 
piicrilis Cameron and S. afghcmica Lelej) inner 
mandibular margin with two small preapical 
denticles (Fig. 9) and gastral tergum 2 with 
large spot of pale pubescence on posterior mar- 
gin (Fig. 4) 

Sc/uanuilotllla sensu Mickel 

4. Apterous; mandible extremely widened apical- 
ly, with deep preapical emargination; hind coxa 
ventrally with small, sharp posterolateral den- 
ticle; gastral sternum 8 membranous laterally, 
narrow median part with parallel lateral ridges 

Spiliinuililla Ashmead 

- Alate; mandible not widened apically, inner 
border with two small preapical denticles and 
larger basal one (cf Fig. 9); hind coxa ventrally 
without denticles; gastral sternum 8 not mem- 
branous laterally, with median carina or tuber- 
cle; ga.stral terga 2-5 very often with median 
longitudinal carina 

Scjiiciinuli>tllUi sensu Mickel 

Bethsmyrmilla Krombein and Lelej, 
new genus 

(Figs. 1, 6, 10-18) 
Type species. — Bethsmynnilla alticola 
Krombein and Lelej, new species. The ge- 
nus is monotypic. 



Female. — Head large, distinctly wider 
than thorax, viewed from above the sides 
gradually widened behind eyes (Fig. 14), 
widened area forming a sharp tubercle as 
viewed from behind (Fig. 15); mandible not 
widened apically, inner margin with sub- 
basal tooth (Fig. 6); medial clypeal lobe 
with prominent lateral tooth (Fig. 10); fla- 
gellomere I 1.8-1.9X its maximal width 
and 1.8-2. Ox as long as flagellomere II 
(Fig. 1 1), the latter wider than its length. 

Mesosoma dorsally more or less quad- 
rangular, laterally crenulated (Fig. 15); me- 
sopleuron with trituberculate supracoxal ca- 
rina (Fig. 16, sc); propodeal dorsum pos- 
teriorly without denticles, posterior face of 
propodeum sloping gently downward; fore 
tarsi with weak comb (Fig. 18); mid coxa 
ventrally with blunt tubercle and mid tro- 
chanter with narrow apical process (Fig. 
12). 

Dorsum of gaster with spots and bands 
of golden pubescence (Fig. 1); basal half of 
gastral sternum 2 with weak median carina 
that does not terminate in a tubercle. 

Male.- — Unknown. 

Discussion. — Differences of this new ge- 
nus from other Oriental Myrmillinae are 
noted in the key. The female of Bethsmyr- 
milla is similar to those of Sqiiamulotilla 
lamellata Mickel and S. amndinacea Pag- 
den in having a lateral tubercle on the head, 
similarly shaped mandibles (cf Figs. 6, 7) 
and similar spots and bands of golden pu- 
bescence (cf Figs. 1, 2) on the gaster It is 
easily separated from the latter species in 
lacking a tubercle on the fore coxa, in hav- 
ing a narrow apical process on the mid tro- 
chanter and the position of the lateral tu- 
bercle on the head, behind the eye in Beths- 
myrmilla and beneath the eye in the latter 
two species. Also, the propodeal dorsum in 
the former is not denticulate posteriorly 
whereas the latter species have at least an 
acute median denticle. 

The female of Bethsmyrmilla is superfi- 
cially similar to another larger group of 
Squamiilotilla species in having spots and 
bands of golden pubescence on some of the 



VOLUME 101. NUMBER 1 



145 







Figs. 1-12. Females of Oriental Myrmillinae. 1-5, Patterns of pale pubescence on abdominal terga. 1, 
Bethsmyrmilla alticola holotype, Sri Lanka, terga 1-3 only. 2, Squamiilotilla lainellata. Vietnam. 3, Spilonnitilla 
consolidata (Cameron), Sri Lanka. 4, Sq. puerilis. South India. 5, Sq. strangulata (Smith), South China. 6-9, 
Mandible. 6, B. alticola. 7, Sq. lamellata. 8, S. consolidata. 9, Sq. puerilis. 10-12, B. alticola. 10. Clypeus, 
frontal view. 1 1, Antennal segments 1-5. 12, Coxa, trochanter, base of femur, mid leg above, hind leg below. 



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Figs. 13-18. BethsDiynnilla alticola. female paratype uncoated. 13, 14, Head. 13, Frontal view (note genal 
tubercle extending laterad of each eye on lower margin). 14, Dorsal view. 15, 16, Thorax. 15, Dorsal view. 16, 
Oblique view, bracket and sc indicate supracoxal carina. 17, Head, posterolateral view, note sparse, long setae 
that margin eye. 18, Foretarsus, note that basal segment is angled, foreshortened. 



VOLUME 101, NUMBER 1 



147 



gastral terga but differs markedly in having 
the lateral tubercle on the head, in lacking 
denticles along the posterior margin of the 
thoracic dorsum and in the gently rather 
than abruptly sloping posterior surface of 
the thorax. 

Etymology. — We take great pleasure in 
naming this remarkable mutillid for Beth B. 
Norden, collector of the type series of B. 
alticola and discoverer of its probable host 
species. 

Bethsmyrmilla alticola Krombein and 
Lelej, new species 

(Figs. 1,6, 10-18) 

Female. — Length (through gastral seg- 
ment 3) 5.0 mm. Red; gaster black except 
sternum 1 totally and tergum 1 basally red; 
mandible red, brownish apically; palpi 
brownish; scape red, flagellomeres red, 
darkened above; legs red, mid and hind tib- 
iae with darker spines; mandible and clyp- 
eus with long pale erect setae, scape with 
shorter ones; frons and vertex with sparse 
recumbent, short reddish setae, genae with 
whitish ones; vertex and frons along eye or- 
bit with sparse, long, erect black setae (Fig. 
17); thoracic dorsum with sparse, short, 
subappressed black setae mixed with long, 
erect black ones; legs with subappressed 
and erect yellowish setae; pattern of golden 
pubescence on gaster (Fig. 1); gastral sterna 

2 and 3 posteriorly with whitish fascia; pro- 
podeal hindface, gastral tergum 1 anterior- 
ly, gastral sternum 2 and gastral terga 2 and 

3 laterally with sparse, erect, whitish setae. 
Head parallel behind eyes, gena postero- 

laterally with strong acute tubercle (Figs. 
13-17); antennal scrobe well developed be- 
tween antennal tubercle and lower part of 
eye (Fig. 13); clypeus delimited above by a 
weak convex carina with four small tuber- 
cles and ending laterally in a strong tooth 
(Figs. 10, 13); antenna with short flagello- 
meres (Fig. 11), flagellomere I 1.8-1.9X its 
apical width and 1.8-2. OX flagellomere II, 
the latter 0.75 X its width; hypostomal ca- 
rina without projection; gena below not car- 
inate; head above with dense large punc- 



tures (Figs. 13, 14, 17). Thorax viewed 
from above (Fig. 15) more or less quadran- 
gular with slightly widened pronotal and 
propodeal areas; humeral angles developed, 
lateral margin of pronotum with vertical 
ridge (Fig. 16); thoracic dorsum noticeably 
convex, weakly serrate laterally, with one 
lateral tubercle on the middle of mesono- 
tum, and with dense reticulate punctures; 
propodeal dorsum posteriorly not dentate 
nor serrate; mesopleuron concave with su- 
pracoxal carina well developed, the latter 
with three blunt tubercles (Fig. 16, sc); mid 
coxa ventrally with blunt tubercle and mid 
trochanter with narrow apical process (Fig. 
12); hind coxa carinate along posterior mar- 
gin of ventral surface, carina ending in a 
blunt tubercle. 

Gastral sternum 1 with well developed 
median carina; gastral tergum 2 with rather 
short lateral felt line and dense small punc- 
tures; gastral sternum 2 with more or less 
flattened central disc and dense punctures 
which are much sparser and larger on disc; 
gastral segments 4-6 lacking in holotype 
and 3-6 in paratype (these parts acciden- 
tally amputated and lost during nest dig- 
ging). 

Male. — Unknown. 

Range. — The species is known only from 
a trail at about 1,950 m altitude along the 
upper border of the Hakgala Botanic Gar- 
den about 10 km S of Nuwara Eliya. We 
believe that it may be more widely distrib- 
uted. Its probable host has been collected 
only at high altitudes, 1,700-1,950 m, at 
various localities in the districts of Nuwara 
Eliya (Nuwara Eliya, Hakgala, Horton 
Plains) and Kandy (Adam's Peak Trail). 

Type material. — Holotype 9, Sri Lanka, 
Nuwara Eliya District, Hakgala Botanical 
Garden, 6°55'N, 80°49'E, 21-22 April 
1997, B.B. Norden [USNM]. Paratype 9, 
same data as holotype but 24-26 February 
1997 [IBPV]. 

Etymology. — The specific name is from 
the Latin altus, high, and -cola, dweller. 

Natural history. — Beth Norden excavated 
two nests of Lasioglossum (Sudila) alphe- 



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mini one each on 26 February and 22 April 
1997. Each nest contained a single female 
mutillid. Both wasps were found at the ba- 
ses of the nest tunnels and are presumed to 
have traveled to the farthest reaches of the 
vertical shafts in an effort to avoid capture. 

The February nest shaft reached a depth 
of 7 cm and also contained a female L. al- 
phenum and a female of the parasitic bee 
Nomada priscilla Nurse that were captured 
a few mm above the mutillid. The April 
nest shaft reached a depth of 13 cm. In this 
nest, two female L. alphenum were collect- 
ed at depths of 12 and 12.5 cm respectively. 

The mutillids moved more vigorously 
than the bees and were more difficult to col- 
lect. Pollen and immature bees (larvae and 
pupae in various stages of development) 
were found in cells located above the base 
of the nest where the wasps were retrieved. 

Discussion. — Norden et al. (1994) found 
two females of the mutillid wasp Pseudo- 
methoca bethae Krombein within a com- 
munal nest of an Arizona bee Exomalopsis 
(Phanamalopsis) solani Cockerell. One 
mutillid was within a provisioned cell pre- 
sumably to feed on the pollen-nectar mass. 
Although neither specimen of B. alticola 
was in a cell with provisions, we suspect 
that they would use this convenient source 
of food while they remained within the 
nest. And when bee brood reached the ap- 
propriate developmental stage it would be 
available for parasitism. Such cryptic be- 
havior by the mutillid would greatly lessen 
exposure to predators and may explain the 
rarity of this species in collections. 

Acknowledgments 
Within the Smithsonian Institution we 
thank Beth B. Norden for her notes on the 



natural history of B. alticola and preparing 
specim.ens for SEM study, Susann G. Bra- 
den for skillful preparation of the scanning 
electron micrographs and George L. Vena- 
ble for preparation of the plates. We are 
grateful to P. Klimov (IBPV) for preparing 
Figures 1-12. ASL thanks the late Borge 
Petersen for generously providing valuable 
exchange material of Oriental mutillid 
wasps and V. Kuznetsov (IBPV) for the gift 
of material. KVK thanks Bryan N. Dan- 
forth, Cornell University, Ithaca, NY, for 
identification of Lasioglosswn (Sudila) al- 
phenum and Maximilian Schwarz, Ansfel- 
den, Austria, for identification of Nomada 
priscilla. We are grateful to G. Nonveiller, 
Zemun, Yugoslavia, for critical reading of 
the manuscript and valuable comments, and 
also an anonymous reviewer 

Literature Cited 

Mickel, C. E. 1933. The Mutillidae of Eastern Asia. 
Lingnan Science Journal 12(3): 289-325. 

. 1935. The Mutillid Wasps of the Islands of 

the Pacific Ocean (Hymenoptera: Mutillidae). 
Transactions of the Royal Entomological Society 
of London 83(2): 177-312. 

Nonveiller, G. 1995. Recherches sur les Mutillides de 
TAfrique XVII. Note pour servir a la connais- 
sance du genre Phstomiitilla Ashmead, 1903 
(male, femelle) avec description du male encore 
inconnu du genre, d'espece nouvelles du genre et 
des nouveaux sous-genres Diaccmthotilla (femel- 
les) et Acanthomutilla (femelles) (Hymenoptera, 
Mutillidae). Entomofauna 16(5): 29-119. 

Norden, B. B., K. V. Krombein, and S. W. T. Batra. 
1994. Nests and Enemies of Exomalopsis (Phan- 
amalopsis) solani Cockerell (Hymenoptera: Apo- 
idea, Mutillidae; Diptera: Asilidae). Proceedings 
of the Entomological Society of Washington 
96(2): 350-356. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 149-163 

TWO NEW WESTERN NEARCTIC CULICOIDES LATREILLE 
(DIPTERA: CERATOPOGONIDAE) DESCRIBED FROM ALL STAGES 

Mark S. Breidenbaugh and Bradley A. Mullens 

Department of Entomology, University of California, Riverside, California 92521, 
U.S.A. (BAM, corresponding author for reprints) 



Abstract. — The egg, larva, pupa, and adult of two new species of Culicoides La- 
treille from desert mountains in southern California and Baja, Mexico are described: 
C. kettlei and C. vetustus. Descriptions of immatures were made from laboratory- 
reared material. Rearing techniques and notes on behavior observed in the laboratory 
are presented. 

Key Words: Culicoides, Ceratopogonidae, immatures, morphology 



Bluetongue and epizootic hemorrhagic 
disease viruses have been suspected of con- 
tributing to lamb mortality in desert bighorn 
sheep populations in southern California 
(DeForge et al. 1982, Wehausen et al. 1987, 
Elliott et al. 1994). A comprehensive sur- 
vey of potential vectors {Culicoides Latre- 
ille) occurring in bighorn habitats within 
the desert mountains of southern California 
(Mullens and Dada 1992a) revealed three 
undescribed species of Culicoides. The 
adults of one species, Culicoides boydi 
Wirth and Mullens have been previously 
described (Wirth and Mullens 1992). The 
present study describes all life-stages of 
these remaining two species and provides 
data on their biology and geographic distri- 
bution. 

Materials and Methods 

Host-seeking females were collected us- 
ing suction traps baited with 1 kg of dry 
ice. Some females collected at the Philip L. 
Boyd Deep Canyon Desert Research Center 
near Palm Desert, Riverside Co., CA, or 
near the town of Morongo Valley, San Ber- 
nardino Co., were given the opportunity to 
feed on heated, defibrinated sheep blood 



through a parafilm or chick-skin membrane 
(Hunt 1994). Engorged females were sep- 
arated from other midges in the collections 
while they were immobilized on a chill ta- 
ble. Female midges were then held at 21°C 
for 7 days. Some individual females would 
oviposit onto damp filter paper, when held 
overnight in a petri dish. Gravid females 
which did not oviposit were decapitated to 
induce oviposition. 

Filter paper with eggs from an individual 
female was placed in a petri dish containing 
nutrient-enriched 1.5% noble agar. Eggs 
were held in a humid chamber and checked 
daily for hatch. First-instar larvae were of- 
fered several food sources, including a nu- 
trient rich liquid diet consisting of bacteria, 
algae, and yeast used for rearing colonized 
Culicoides variipennis sonorensis Wirth 
and Jones (Jones et al. 1969). In addition, 
the bacterial feeding nematodes, Pelodera 
sp. and Panagrellus redivivus (L.), were 
supplied as potential prey on a biweekly ba- 
sis (Mullens and Velten 1994). Larval 
growth and feeding behavior were observed 
daily and the duration of egg, larval, and 
pupal development recorded. 

Samples of eggs from associated females 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



were placed into 70% EtOH. Specimens 
were fixed (Day et al. 1997), then critical- 
point dried, transferred onto stubs backed 
with sticky tape, sputter-coated with gold- 
palladium, and viewed on either a JOEL 
JSM-35C or Phillips XL30 scanning elec- 
tron microscope. 

The parental adult female was preserved 
in 70% EtOH and slide-mounted in balsam 
after Wirth and Marston (1968). Fourth-in- 
star larvae, pupae, and adults from reared 
offspring of the parental female were also 
preserved. Offspring were mounted in Can- 
ada balsam or Hoyer's medium. Descrip- 
tions of immature stages and males of both 
species were made from laboratory-reared 
material. 

The terminology of Downes and Wirth 
(1981) is used for adults, of Lamberson et 
al. (1992) and Nevill and Dyce (1994) for 
pupae, of Murphree and Mullen (1991) for 
larvae, and of Becker (1961) and Campbell 
and Kettle (1975) for eggs. Nomenclature 
agrees with Borkent and Wirth (1997) and 
Spinelli and Ronderos (1997). 

The following measurements were made 
from fourth-instar larvae: total length (TL), 
head length (HL), head width (HW), sub- 
genal width (SGW), mandible length (ML), 
width across the lateral arms of the epi- 
pharynx (LAW), total width across the 
paired dorsal comb sclerites of the epiphar- 
ynx (DCW), caudal-segment length (CSL), 
caudal-segment width (CSW), length of se- 
tae 'o' (OL), and the distance between their 
bases. 

From values listed above, the following 
ratios were calculated: head ratio (HR = 
HL/HW), subgenal ratio (SGR = HW/ 
SGW), and caudal-segment ratio (CSR = 
CSL/CSW). Illustrations were made of the 
morphology and chaetotaxy of the head 
capsule and caudal segment. The hyposto- 
ma, epipharynx, hypopharynx, and mandi- 
ble were illustrated, and thoracic pigmen- 
tation and anal papillae were drawn when 
appropriate. 

For pupae the following structures were 
described and illustrated: respiratory horn. 



operculum, caudal segment, and the ad, dl, 
dasm, dpm, lasm, Ipm, and vpm tubercles. 
The holotype, allotype, and some para- 
types are deposited in the National Museum 
of Natural History (USNM), Smithsonian 
Institution, Washington, DC; paratypes, as 
available, will be deposited in the collec- 
tions of the University of California, Riv- 
erside and the California Academy of Sci- 
ences, San Francisco. 

Ciilicoides (Haematomyidium) kettlei 
Breidenbaugh and Mullens, new species 

(Figs. 1-4) 

Egg. — Banana-shaped. Surface with two 
types of ansulae arranged in longitudinal 
rows (Fig. lA). Moderately stalked ansulae 
present on concave surface, ridges com- 
posed of flattened papillate ansulae on con- 
vex surface (Fig. IB). Average length = 
356 ± 17 |x; width = 53 ± 2 |x (n = 11). 

Larva.— Total length = 2.79 (2.16-3.51, 
n = 15) mm. Head capsule (Figs. 2A-C): 
Light brown. Small larvae, HL = 128 (122- 
134, n = 29) fx, HW = 89 (82-102, n = 
28) |jl; shape somewhat long and narrow, 
SGW = 58 (54-67, n = 30) |jl; HR = 1.4 
(1.2-1.6, n = 27); SGR = 1.5 (1.4-1.8, n 
= 27). Mandible (Fig. 2F) medium-small, 
ML = 35 (22-38, n = 29) |x; with pointed 
apex and subapical rounded prominence; 
base with distinct point of articulation. Hy- 
postoma (Fig. 2G) rounded, entire. Labium 
small, crescent-shaped and sclerotized. Epi- 
pharymx (Fig. 2H): Dorsal-comb sclerites 
with 5-7 unequal angular teeth; moderately- 
wide, DCW = 11 (10-13, n = 17) |jl. Comb 
2 narrow with short, rounded teeth; comb 4 
wider, with rounded teeth. LAW = 49 (43- 
52, n = 30) |x; lateral curtains composed of 
short, dense, filamentous processes. Hypo- 
pharynx apparently poorly sclerotized, not 
seen. Thoracic pigmentation (Fig. 21): dis- 
tinct lateral pattern on thoracic segments. 
Prothorax with arched pattern; mesothorax 
and metathorax with crescent and sagittate 
patterns, respectively. Caudal segment (Figs. 
2D, E): Short, CSL = 231 (189-285, n = 
32) |JL and moderately- wide, CSW = 126 



VOLUME 101. NUMBER 1 



151 




Fig. 1. Culicoides kettlei, egg. A, Lateral view. B, Detail of surface. 



(90-179, n = 31) |x; Oval; CSR = 1.9 (1.5- 
2.2, n == 31). Setae "o" short relative to the 
CSL, OL - 46 (38-51, n = 19) fx; bases of 



setae moderately separated, OD = 37 (26- 

45, n = 19) |JL. Anal papillae not observed. 

Material studied. — Laboratory-reared from 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Ciilicoides kettlei, larva. A-C, Head capsule. A, Dorsal view. B, Ventral view. C, Lateral view. D, 
Caudal segment, dorsal view. E, Caudal segment, ventral view. F, Mandible. G, Hypostoma. H, Epipharynx. I, 
Thoracic pigmentation, dorsal view. 



eggs deposited by individual females collect- 
ed in San Bernardino County, CA, Morongo 
Valley, 15-VI-95 (n = 3); 22-V-96 (n = 5); 
14-VI-96 (n = 24), paratypes: slides 
25,26,27. 

Pupa. — Light yellow brown. Respiratory 
horn (Fig. 3A): yellow brown; basal half 
crenulated; spines, sparsely distributed. 
Four lateral spiracules present and 8-1 1 
apical spiracles. Tracheal annulations asso- 
ciated with tracheae visible in basal portion 
and a reticulated pattern is visible in distal 
portion. Operculum (Fig. 3B): Yellow 
brown, with two types of spines, neither ex- 
tending beyond the am tubercles. Sagittate 
spines occurring laterally; smaller, rounded 



spines occurring medially. Anterior margin 
moderately notched. Caudal segment (Fig. 
3C): Spines present at base of posterolat- 
eral processes, absent apically; band of 
spines along anterior margin and a patch of 
spines located basimedially on dorsum. 

Chaetotaxy: Dorsal tubercles (Fig. 3D): 
1 and 2 round with stout spine; 3 round 
with a short seta; 4 a slender seta; 5 a small, 
round pore, ad tubercle (Fig. 3E): spinate 
with two subequal spines, dl tubercle (Fig. 
3F): with 3 unequal, slender spines. Abdo- 
men (Fig. 3G): dasm tubercles: 1 with a 
spine and 2 a bristle, dpm tubercles: round- 
ed; 1 and 2 with short spine; 3 and 4 lack 
setae; 5 with a short hair lasm tubercle: spi- 



VOLUME 101. NUMBER 1 



153 




y^hj K=^ '^ 



Fig. 3. Culicoides kettlei, pupa. A, Respiratory horn. B, Operculum. C, Caudal segment. D, Dorsal tubercles. 
E, ad tubercle. F, dl tubercle. G, Abdominal chaetotaxy. 



nate with short spine. Ipm tubercles: 1-3 
spinate; 1 and 3 with short spine; 2 with a 
bristle, vpm tubercles: rounded; 1 with a 
spine; 2 and 3 with long and medium length 
setae, respectively. 

Material studied. — Laboratory -reared 
from individual females collected in San 
Bernardino County, CA, Morongo Valley, 
14-V-95 (n = 6), paratypes: slides 10,11. 

Adults. — Female: Wing length 1.06 
(0.92-1.19, n = 28) mm. Head: Eyes (Fig. 
4A) bare; separated by a single facet width; 
without interfacetal hairs. Antenna (Fig. 



4B) with lengths of flagellomeres of holo- 
type 43-27-28-33-29-3 1 -30-33-36-37-43- 
47-63 (in p.); antennal ratio 0.91 (0.74- 
1.04, n = 28); well-developed sensilla coe- 
loconica present on flagellomeres 1, 4-8; 
26% (7/27) of observed specimens lack 
sensilla on flagellomere 4. Palpus (Fig. 4C) 
with lengths of segments 24-46-61-20-26; 
palpal ratio 2.07 (1.75-2.36, n = 29); P/H 
ratio 0.79 (0.70-0.86, n = 18); third seg- 
ment swollen with broad shallow sensory 
pit. Thorax: Brown, scutum without con- 
spicuous pattern. Legs brown (Fig. 4D), 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 











''% 



^^., 



''^>^r. 



\ 



TjT(]]-rn]7rT7-i/77v^-|-ri-fnV^'n''''"^'^^ 



H'n'^ 



. - - - -^;^ 
- - " , - <> 

^^ E 



Fig. 4. Culicoides kettlei, adult. A-F Female, G-H, Male. A. Eye separation. B. Antenna. C, Palpus. D, 
Legs, left to right, front, mid. and hind. E, Wing. F Spermathecae. G. Genitalia, parameres omitted. H, Parameres. 



femorotibial joints blackish; fore and mid 
femora with narrow subapical pale rings, all 
tibiae with narrow subbasal pale rings and 
hind tibiae with distal fourth pale; hind tib- 
ial comb with five spines, second from the 
spur longest. Wing (Fig. 4E) with a slightly 



angular poststigmatic pale spot; distinct 
transverse pale spot present in r5; cell ml 
with narrow pale spot; cell m2 with round- 
ed pale spot located distally and with a lon- 
gitudinal pale spot lying posterior to medial 
fork. Macrotrichia in moderate numbers 



VOLUME 101, NUMBER 1 



155 



evenly distributed over wing. Halter pale. 
Abdomen: Brown. Two slightly unequal 
spermathecae, plus rudimentary third and 
sclerotized ring (Fig. 4F). Functional sper- 
mathecae with long slender necks; length, 
including neck, 32 (29-37, n = 25) |jl for 
the larger, 29 (26-35, n = 26) |x for the 
smaller. 

Male: Genitalia (Figs. 4G-H): Ninth 
sternum with scarcely perceptible caudo- 
median excavation, ventral membrane spic- 
ulate; apicolateral processes moderately 
long, pointed, caudal margin between them 
slightly cleft medially. Gonocoxite moder- 
ately stout; ventral root with anterior point 
longer, more slender than dorsal root; gon- 
ostylus moderately curved and slender with 
moderately bent, pointed tip. Aedeagus 
with basal arch extending more than two- 
thirds of total length; distal portion short, 
tapering to a simple tip. Paramere (Fig. 4H) 
with strong basal knob; midsection arched; 
distally, greatly narrowed and reflexed upon 
midsection, constricting to a noticeably 
fringed filamentous tip. 

Distribution. — Southern California and 
northern Baja California, Mexico. 

Material studied.— HOLOTYPE: $ , Riv- 
erside County, CA, Deep Canyon, Bighorn 
Overlook, nr. Palm Desert, 22-IX-88 (B. A. 
Mullens), CO.-baited trap. ALLOTYPE: S, 
San Bernardino County, CA, Big Morongo 
Canyon, Morongo Valley, 15-VI-95 (Brei- 
denbaugh), laboratory-reared from wild- 
collected female. PARATYPES: CALI- 
FORNIA. 6 9, same data as holotype, ex- 
cept 1 $ 24 May 1988; 6 $, Riverside Co., 
Bighorn Drive, Palm Desert, 8 October 
1988, CO.-baited trap, 1 9, same except 22 
September 1988; 9 9,4 6, same data as 
allotype except from June 1995 to July 
1996 (some K. Luhring); MEXICO. 7 9, 
Mexico, Baja California, Cadavina, 19 
March 1994 (C. Szijj), CO.-baited trap. 

Behavior and rearing. — In the laboratory, 
the average female fecundity was 74 ± 24 
eggs (n = 10). From a single reared cohort 
observed daily, pupation first occurred 26 
days following egg hatch. In culture, larvae 



fed on Pelodera sp. nematodes. Adults have 
been collected from soil-emergence traps 
along a seasonal creek in the Deep Canyon 
watershed (Breidenbaugh and Mullens, in 
preparation), indicating that creek margins 
are used by this species as a larval devel- 
opment site. Feeding behavior is poorly 
known with a single report of an unfed fe- 
male recovered from a bighorn sheep (Mul- 
lens and Dada 1992b). 

Discussion. — Culicoides kettlei, was 
originally recognized by the late W. W. 
Wirth who referred to this species as #120 
(W. W. Wirth, personal communication). 
Following Vargas (1960) the male genitalia 
fit nicely into the subgenus Haematomyi- 
dium. The female wing pattern is inconclu- 
sive, since the r-m crossvein is not dark. 
However, we feel that placement in the sub- 
genus Haematomyidium is reasonable. 
Adult morphology is similar to that of Cu- 
licoides {Haematomyidium) debilipalpis 
Lutz, a common biting midge in the eastern 
U.S.. From 1985-1997 C. debilipalpis was 
synomized with C. lahillei Iches (Spinelli 
and Wirth 1985). Recently, however, the 
validity of C. debilipalpis was confirmed 
(Spinelli and Ronderos 1997). The range of 
C debilipalpis includes the southeastern 
U.S. as far west as Louisiana, with a dis- 
junct distribution that includes Costa Rica 
south to Argentina. In contrast, C. lahillei 
is strictly South American. The females of 
C. kettlei can be easily separated from C. 
debilipalpis and all other Nearctic members 
of Haematomyidium by the antennal sen- 
sorial pattern (l,4(5)-8) (Wirth et al. 1985). 
In addition, no Neotropical species of Hae- 
matomyidium are known to occur north of 
southern Mexico (Wirth et al. 1988), and 
their range thus does not overlap with that 
of C. kettlei. 

Apart from the similarity in size, the lar- 
vae of this species are noticeably different 
from those of the only other North Ameri- 
can species in this subgenus with described 
larvae, Culicoides paracusis (Goel- 
di)(Murphree and Mullen 1991). The hy- 
postoma is round and smooth in this species 



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while lateral teeth are present in C. paraen- 
sis. Furthermore, the hypostoma lacks the 
distinct subapical notch seen in C. paraen- 
sis. The pupa of this species is similar to C. 
paraensis in the type and location of spines 
on the operculum. 

Etymology. — This species is named in 
honor of Dr. D. S. Kettle, Emeritus Profes- 
sor, Department of Entomology, University 
of Queensland, Australia, for his pioneering 
work on the biology of CuUcoides, specif- 
ically in the discipline of the morphology 
of immatures. 

CuUcoides vetustus Breidenbaugh and 
Mullens, new species 

(Figs. 5-8) 

Egg. — Cigar-shaped. Surface with flat- 
tened longitudinal rows, many not contig- 
uous from end to end (Fig. 5A). Ansulae 
flattened, not distinct, merge to form lon- 
gitudinal ridges (Fig. 5B), present on all lat- 
eral surfaces, and not varying with curva- 
ture. Average length = 267 ± 28 |x and 
width = 49 ± 3 |JL (n = 8). 

Larva.— Total length = 3.25 (2.81-4.0, n 
= 15) mm. Head capsule (Figs. 6A-C): 
Yellow; medium sized, HL = 180 (166- 
192, n = 28) |x, HW = 123 (109-138, n = 
27) |JL, SGW = 90 (80-102, n = 28) jx; 
overall shape long and narrow, HR = 1.5 
(1.2-1.7, n = 27), very oblong, SGR = 1.4 
(1.2-1.5, n = 27). Mandible (Fig. 6D) me- 
dium length, ML = 5 1 (48-54, n = 24) |x; 
base wide; pointed marginal prominence 
basimedially, followed by a subapical 
notch, sharply angled to pointed apex. Hy- 
postoma (Fig. 6E) difficult to see but round- 
ed medially and smooth. Epipharynx (Fig. 
6F): Dorsal-comb sclerites moderately 
wide, DCW = 14 (13-15, n = 24) |jl, with 
5 subequal pointed teeth/sclerite; comb 4 
with many unequal pointed, rounded teeth; 
lateral curtains wide; teeth thin, hair-like; 
LAW wide relative to DCW, LAW = 60 
(52-67, n = 21) jx, indented near lateral 
apex. Hypopharynx (Fig. 6G) with hypo- 
pharyngeal fringe separated into two dis- 
tinct prominences by a medial notch. Tho- 



racic pigmentation (Fig. 6H): Absent. Cau- 
dal segment: Short with length varying 
considerably among individuals, CSL = 
286 (131-326, n = 25) fx; narrow, CSW = 
139 (86-157, n = 26) jx, CSR = 2.1 (1.5- 
2.4, n = 25); setae "o" of medium length, 
OL = 82 (61-99, n = 18) |x, and well sep- 
arated, OD = 45 (35-53, n = 21) |x. Anal 
papillae (Fig. 61) four deeply bifurcate 
pairs. 

Material studied. — Laboratory-reared 
from eggs deposited by individual females 
collected in San Bernardino County, CA, 
Morongo Valley, 15-VI-95 (n = 6); 14- V- 
96 (n = 15); 14-V-96 A42 (n = 15), para- 
types: slides 2-7. 

Pupa. — Light to dark brown. Respiratoty 
horn (Fig. 7A): Proximal portion light 
brown; roughly divided into 3 subequal 
portions. First section with annular trache- 
ae; second portion annulations obscured by 
reticulation pattern; 3 lateral spiracular pro- 
tuberances present; terminal section, dark 
brown with 5-9 spiracular openings apical- 
ly; few scales on horn. Operculum (Fig. 
7B): Mostly smooth, 3-8 large spines on 
lateral margins and patch of setae on central 
portion; no large spines medial to the am 
tubercles; large spines triangular. Caudal 
segment (Figs. 7C-D): Narrow V-shaped 
cluster of small spines on dorsal surface of 
both sexes. Anterior band of spines com- 
plete on both sexes; large spines on proxi- 
mal % of posterolateral processes; distal 
third, smooth dark brown. 

Chaetota.xy: Dorsal tubercles (Fig. 7E): 
setae 1 and 2 of medium length, stout; seta 
3 short, stout; seta 4 a long, slender bristle; 
5 a circular pore, ad tubercle (Fig. 7F) with 
one long, one shorter setae, dl tubercle (Fig. 
7G) with a long slender bristle and second 
shorter thicker bristle. Abdomen (Fig. 7H): 
dasm tubercles: setae 1 and 2 of medium 
thickness, 1 longer than 2. dpm tubercles: 
setae 1 and 2 equal; 3 and 4 lack setae; 5 
short, lasm tubercle: rounded with short 
spine. Ipm tubercles: spinate with shorter, 
stouter, equal bristles on 1 and 3, bristle 2 
longer and slender, vpm tubercles: 2 with a 



VOLUME 101. NUMBER 1 



157 




Fig. 5. Ciilicoides vetiistiis, egg. A. Lateral view of entire egg. B, Detail of egg surface. 



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Fig. 6. Culicoides vetustus. larva. A-C, Head capsule. A, Dorsal view. B, Ventral view. C, Lateral view. D, 
Mandible. E, Hypostoma. F. Epipharynx. G, Hypopharynx. H, Head and thorax, dorsal view. I, Anal papillae, 
caudal segment. 



slender bristle, longer and thinner than 1 
and 3. 

Material examined. — Laboratory-reared 
from eggs deposited by individual females 
collected in San Bernardino County, CA, 
Morongo Valley, 14-V-96 (n = 8); 15-VI- 
95 A5 (n = 17), paratypes: slides 17-18; 
14-V-96 A42 (n = 3 ), paratypes: slide 1. 

Adults.— Female: Wing length 1.24 ( 1.1- 
1.4, n = 20) mm. Head: Eyes (Fig. 8A) 
separated by single facet width; without in- 
terfacetal hairs. Antenna (Fig. 8B) lengths 



of flagellomeres of holotype 46-28-28-30- 
30-30-33-33-43-46-46-50-65; antennal ratio 
0.97 (0.92-1.0, n = 18); well-developed 
sensilla coeloconica present on flagello- 
meres 1, 11-13. Palpus (Fig. 8C) with 
lengths of segments 24-50-78-30-33; third 
segment swollen, with round moderately 
deep sensory pit near apex. Palpal ratio 
2.06 (1.8-2.3, n = 21); proboscis long, P/ 
H ratio 0.95 (0.81-1.3, n = 19). Thorax: 
Brown, lateral aspects of scutum darker. 
Legs brown, lacking pattern; hind tibial 



VOLUME 101. NUMBER 



159 




Fig. 7. Ciilicoides vetustus. pupa. A, Respiratory horn. B, Operculum. C, Caudal segment, male. D, Caudal 
segment, female, E, Dorsal tubercles. F, ad tubercle. G, dl tubercle. H, Abdominal chaetotaxy. 



comb (Fig. 8D) with 5 spines, second from 
spur longest. Wing (Fig. 8E) uniformly 
grayish with pale spot on anterior margin 
just past second radial cell. Halter pale. Ab- 
domen: Brown. Two slightly unequal sper- 
mathecae (Fig. 8F), plus rudimentary third; 
functional spermathecae with short slender 
necks; length 44 (35-51, n = 18) fx for larg- 
er, 42 (34-50, n = 17) |jl for smaller. 

Male: Genitalia (Figs. 8G-H): Sternite 9 
with broad moderately deep caudomedian 
excavation, ventral membrane spiculate; 
tergite 9 with prominent triangular apico- 
lateral processes, moderately broad, slightly 
divergent; the caudal margin between them 
transverse without medial cleft. Gonocoxite 
moderately stout; ventral root more slender 



than dorsal root; gonostylus curved distally, 
moderately slender, with moderately broad 
blunt tip. Aedeagus Y-shaped; basal arch 
extending half of total length; basal arms 
short, moderately slender, recurved at ex- 
treme apex, distal portion moderately 
broad. Paramere (Fig. 8H) with distinct an- 
terior process; midpoint straight; apex di- 
verging with three sharp medial spines, tip 
sharply pointed, diverging. 

Distribution. — Mojave and Colorado de- 
serts of southern California. 

Material examined. — HOLOTYPE: 9, 
Riverside County, CA, Deep Canyon, nr 
Palm Desert, 29-VI-89 (B. A. Mullens), 
COj-baited trap. Allotype: (5, San Bernar- 
dino County, CA, Big Morongo Canyon, 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



8 





- - ^ 



x 



"n>i„ 




'')'^^''Tl'i\ll^l|'llrpyl^^^^^TV^^t^''•T'^'"^'^'* 



H-llW 



,-.Af 



■^^^ 





Fig. 8. Culicoides vetiistus. adult. A-F, Female. G-H, Male. A. Eye separation. B, Antenna. C, Palpus. D, 
Hind leg. E, Wing. F Spemiathecae. G. Genitalia, parameres omitted. H, Parameres. 



Morongo Valley, 15-VI-95 (Breidenbaugh), 
laboratory-reared from wild-collected fe- 
male. PARATYPES: 1 9, same data as al- 
lotype; 16 $, 11 6 same data as allotype, 
except collected between 14-V-96 and 2- 
VII-96, some reared from wild-collected 9 ; 
9 9 , San Diego County, CA, Anza Borrego 
State Park, Yaqui Well, 26-1 V-97 (Breiden- 
baugh), C02-baited trap. 

Biology and rearing. — Adult females 
collected in C02-baited traps readily fed 



through an artificial membrane and depos- 
ited eggs in the laboratory. Larvae exhib- 
ited the slowest development of any species 
reared on the agar system. Average clutch 
size was 86 ± 35 eggs (n = 52 females); 
these hatched in 7 ± 1 days with 67 ± 25% 
fertility (n = 25 females). Average devel- 
opment period to pupation was 117 ± 28 
days (n = 46). Adults emerged 4 ± 1 days 
later (n = 39). 

It is not known if the rearing medium 



VOLUME 101. NUMBER 1 



161 



was responsible for the slow development 
or whether this is the normal condition for 
larvae of this species. The percentage of 
larvae reaching adulthood (approx. 31%) 
was similar to the other species reared 
(Breidenbaugh and Mullens, in prepara- 
tion). Despite extensive observations, lar- 
vae were never seen feeding on nematodes; 
thus, this species was probably utilizing 
other microorganisms present in the cul- 
tures. 

Discussion. — As discussed by Blanton 
and Wirth (1979), adults of a number of 
Nearctic CuUcoides do not key readily to 
the subgenera described by Vargas (1960). 
This is true for C vetustus. From adult and 
larval characteristics presented here, we are 
tentatively placing C vetustus in the bigut- 
tatus species group. For example, the man- 
dibles of the larvae resemble those illus- 
trated by Murphree and Mullen (1991) of 
CuUcoides biguttatus (Coquillett) and Cu- 
Ucoides spinosus Root and Hoffman except 
for the depth of the notch on the subapical 
margin of the mandible of C vetustus. The 
number of dorsal-comb teeth (5) on the epi- 
pharynx is the same for C. spinosus. How- 
ever, the lateral arms of the epipharynx are 
notched in C vetustus, but not in C spi- 
nosus or C. biguttatus. The hypostoma of 
C. biguttatus and C vetustus are very sim- 
ilar. 

The pupal operculum of C vetustus re- 
sembles that of C biguttatus and C. spi- 
nosus. However, the operculum of C. ve- 
tustus has fewer spines, and these are lim- 
ited to the lateral margins. The respiratory 
horn of C vetustus is more similar to that 
of C. spinosus, darkened at the tip and with 
3 spiracular openings visible. 

The range of C vetustus overlaps only 
with C sublettei Atchley and C. usingeri 
Wirth of the biguttatus group. The male 
genitalia of these species are similar in sev- 
eral respects, including the general shape of 
the parameres. 

The apices of the parameres of C. vetus- 
tus are divided into 3-4 short spines and 
the distomedian process of the aedeagus is 



bluntly rounded. The aedeagus of C. vetus- 
tus is more slender than that of C. sublettei 
or C usingeri, and there are 5 tibial spines 
in C. vetustus and C. sublettei, but 4 in C. 
usingeri. The females of C. vetustus lack a 
very distinctive wing pattern. However, 
they can be distinguished clearly from the 
other species in the biguttatus group and 
from CuUcoides piliferus Root and Hoff- 
man, which it superficially resembles, by 
the sensorial pattern (1, 11-13). 

Etymology. — Latin vetustus for long- 
lived, referring to the lengthy development 
period of the larvae. 

Discussion 

Most descriptions of CuUcoides species 
have included only the adult stage or in 
some cases, only the female. This is a result 
of the relative difficulty in locating devel- 
opmental sites or collecting and associating 
males. The collection of host-seeking fe- 
males, use of an artificial host, and a labo- 
ratory rearing system proved a very suc- 
cessful way of associating immature stages 
with adults. This approach should be useful 
for many species whose adults are known. 
The rearing technique is advantageous in 
that the entire cohort from an isofemale is 
unquestionably conspecific. In contrast, 
field-collected larvae, if they are located, 
generally must be killed and slide-mounted 
for identification. Consequently, an incor- 
rect association with other larvae in the col- 
lection which have been reared to adults is 
possible. 

The rearing method used here probably 
results in a narrower range of measurements 
for morphometric analysis, than if field-col- 
lected material were examined. Environ- 
mental homogeneity and the genetic simi- 
larity of laboratory-reared sibling larvae 
can result in morphological measurements 
that are unnaturally similar. The descrip- 
tions herein, however, will allow field-col- 
lected immatures to be identified and mea- 
sured to better characterize the range of nat- 
ural variability in characters such as size. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Acknowledgments 

We thank K. Luhring (University of Cal- 
ifornia, Riverside) for help with collections 
and rearing; W. W. Wirth (deceased) for as- 
sistance with initial discovery of these new 
species; W. L. Grogan (Salisbury State Uni- 
versity, MD); and the California Academy 
of Sciences which supplied adults of C. de- 
bilipalpis. 

We appreciate the comments of W. L. 
Grogan and C. S. Murphree (Belmont Uni- 
versity, TN) on the manuscript as well as 
A. Borkent (Enderby, British Columbia), 
W. L. Kramer (Nebraska Dept. of Health 
and Human Services, NE), and G. R. Spi- 
nelli (Museo La Plata, Argentina) for ad- 
vice. This study was supported in part by 
the Anderson Endowed Fellowship award- 
ed to MSB and by USDA-NRICGP #94- 
37312-06232 to BAM. 

Literature Cited 

Becker, P. 196L Observations on the life cycle and 
immature stages of Ciilicoides circumscriptiis 
Kieff. (Diptera: Ceratopogonidae). Proceedings of 
the Royal Society Edinburgh 67: 363-388. 

Blanton. E S. and W. W. Wirth. 1979. The sand flies 
{CuUcoides) of Florida (Diptera: Ceratopogoni- 
dae). Arthropods of Florida and Neighboring Land 
Areas, Vol. 10, 204 pp. 

Borkent, A. and W. W. Wirth. 1997. The World species 
of Ceratopogonidae (Diptera). Bulletin of the 
American Museum of Natural History 233, 257 
pp. 

Campbell. M. M. and D. S. Kettle. 1975. Oogenesis in 
CuUcoides brevitarsis Kieffer (Diptera: Cerato- 
pogonidae) and the development of a plastron-like 
layer on the egg. Australian Journal of Zoology 
23: 203-218. 

Day, J., D. Duzak, Y. Braverman, and A. Chizov-Ginz- 
burg. 1997. Ultrastructure of the eggs of CuUcoi- 
des circumscriptus. CuUcoides gejgelensis. and 
CuUcoides imicola (Diptera: Ceratopogonidae). 
Journal of the American Mosquito Control Asso- 
ciation 13: 76-83. 

Deforge, J. R. and J. E. Scott. 1982. Ecological inves- 
tigations into high lamb mortality. Transactions of 
the Desert Bighorn Council 26: 65-76. 

Downes, J. A. and W. W. Wirth. 1981. Chapter 28. 
Ceratopogonidae. pp. 393-421. /// McAlpine, J. F. 
et al., eds. Manual of Nearctic Diptera, Vol. 1. 
Research Branch Agriculture Canada Monograph 
27. 

Elliott, L. E. W. M. Boyce, R. K. Clark, and D. A. 



Jessup. 1994. Geographic analysis of pathogen ex- 
posure in bighorn sheep {Ovis canadensis). Jour- 
nal of Wildlife Diseases 30: 315-318. 

Hunt. G. J. 1994. A procedural manual for the large- 
scale rearing of the biting midge, CuUcoides var- 
iipennis (Diptera: Ceratopogonidae). United States 
Department of Agriculture Research Service, 
ARS-121. 

Jones, R. H., H. W. Potter, and S. K. Baker. 1969. An 
improved larval medium for colonized CuUcoides 
variipcnnis (Diptera: Ceratopogonidae). Journal of 
Economic Entomology 62: 1483-1486. 

Lamberson, C, C. D. Pappas, and L. G. Pappas. 1992. 
Pupal taxonomy of the tree-hole CuUcoides (Cer- 
atopogonidae: Diptera) in eastern North America. 
Annals of the Entomological Society of America 
85: 111-120. 

Mullens, B. A. and C. E. Dada. 1992a. Spatial and 
seasonal distribution of potential vectors of hem- 
orrhagic disease viruses to peninsular bighorn 
sheep in the Santa Rosa Mountains of southern 
California. Journal of Wildlife Diseases 28: 192- 
205. 

. 1992b. Insects feeding on desert bighorn 

sheep, domestic rabbits, and Japanese quail in the 
Santa Rosa Mountains of southern California. 
Journal of Wildlife Diseases 28: 476-480. 

Mullens, B. A. and R. K. Velten. 1994. Rearing Cu- 
Ucoides variipennis sonorensis (Diptera: Cerato- 
pogonidae) on agar and nematodes. Journal of 
Medical Entomology 31: 175-177. 

Murphree, C. S. and G. R. Mullen. 1991. Comparative 
larval morphology of the genus CuUcoides La- 
treille (Diptera: Ceratopogonidae) in North Amer- 
ica with a key to species. Bulletin of the Society 
of Vector Ecology 16: 269-399. 

Nevill, H. and A. L. Dyce. 1994. Afrotropical CuU- 
coides: Description and comparison of the pupae 
of seven species of the simiUs supergroup (Cera- 
topogonidae: Diptera). Onderstepoort Journal of 
Veterinary Research. 61: 85-106. 

Spinelli. G. R. and W. W. Wirth. 1985. Clave para la 
identificacion de las especies del genero CuUcoi- 
des Latreille presentes al Sur de la Cuenca Ama- 
zonica. Nuevas citas y notas sinoni'micas (Diptera: 
Ceratopogonidae). Review of the Society of En- 
tomology, Argentina 44: 49-73. 

Spinelli, G. R. and M. M. Ronderos. 1997. Description 
of the male of CuUcoides lahiUei and revalidation 
of CuUcoides debiUpalpis (Diptera: Ceratopogon- 
idae). Transactions of the American Entomologi- 
cal Society 123: 303-309. 

Vargas, L. I960. The subgenera of CuUcoides of the 
Americas (Diptera, Ceratopogonidae). Revista de 
Biologia Tropical 8: 35-47. 

Wehausen, J. D.. V. C. Bleich. B. Blong, and T L. 
Russi. 1987. Recruitment dynamics in a southern 



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California sheep population. Journal of Wildlife 
Management 51:86-98. 

Wirth, W. W. and N. Marston. 1968. A method for 
mounting small insects on microscope slides in 
Canada balsam. Annals of the Entomological So- 
ciety of America 61: 783-784. 

Wirth, W W., A. L. Dyce, and B. V. Peterson. 1985. 
An atlas of wing photographs, with a summary of 
the numerical characters of the Nearctic species of 
Culicoides (Diptera: Ceratopogonidae). Contribu- 
tions of the American Entomological Institute 22: 
1-46. 



Wirth. W. W., A. L. Dyce, and G. R. Spinelli. 1988. 
An atlas of wing photographs, with a summary of 
the numerical characters of the Neotropical spe- 
cies of Culicoides (Diptera: Ceratopogonidae). 
Contributions of the American Entomological In- 
stitute 25: 1-72. 

Wirth, W. W and B. A. Mullens. 1992. Culicoides boy- 
di (Ceratopogonidae: Diptera): A potential vector 
of hemorrhagic disease viruses to desert bighorn 
sheep in southern California. Journal of Medical 
Entomology 29: 1006-1010. 



PROC. ENTOMOL. SOC. WASH. 

lOUl), 1999. pp. 164-174 

BLASTOBASIS GRAMINEA, NEW SPECIES (LEPIDOPTERA: 

GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASINAE), A STEM BORER 

OF SUGAR CANE IN COLOMBIA AND VENEZUELA 

David Adamski 

Research Associate, Department of Entomology, MRC-127, Smithsonian Institution, 
Washington, DC 2056-0127, U.S.A. 



Abstract. — Blastobasis graminea, new species, a stem borer of sugar cane in Colombia 
and Venezuela, is described and illustrated. For the first time, a larva of Blastibasini is 
described in detail. Scanning electron micrographs of the larva, illustrations of the larval 
mandible, setal maps, and photographs of larval damage and pupation sites are provided. 
Auximobasis obstricta Meyrick 1918, is transferred to Blastobasis Zeller 1855, n. comb., 
and Blastobasis siibolivacea Walsingham 1897, is transferred to Holcocera Clemens 1863, 
n. comb. 

Key Words: Coleophoridae, Blastobasinae, Blastobasis, sugar cane, Colombia, Venezuela 



For decades, entomologists have known 
that larvae of at least one species of micro- 
lepidoptera other than Diatraea saccharalis 
(Fabricius) (Crambidae) feed on sugar cane 
and related grasses in Latin America. Al- 
though adult specimens of one species of 
Coleophoridae (Blastobasinae) have been 
collected since the late 1940's and 1950's 
by H. E. Box in Venezuela and during the 
early 1970's and 1980's by L. Cardenas and 
others in Colombia, this moth remained 
nameless. 

Because many Blastobasinae are similar 
in wing pattern, they are frequently mis- 
identified. For example, the species de- 
scribed herein, Blastobasis graminea, had 
been previously misidentified as Auximo- 
basis obstricta Meyrick 1918, (Box 1953, 
Guagliumi 1962) and Blastobasis siiboli- 
vacea Walsingham 1897, (Martorell 1976). 
In addition, type specimens of Neotropical 
Blastobasinae have not been studied sys- 
tematically until recently. 

Since Meyrick (1894) the Blastobasinae 
have long been considered to be monophy- 



letic; recent studies (Adamski and Brown 
1989, Hodges, in press) have corroborated 
this notion and postulated phylogenetic re- 
lationships of the Blastobasinae within Ge- 
lechioidea. In this study, the Blastobasidae 
(sensii Adamski and Brown 1989) are treat- 
ed as a subfamily within the Coleophoridae, 
following Hodges (in press). 

The purpose of this paper is to describe 
and illustrate Blastobasis graminea, new 
species, and to make available to entomol- 
ogists and sugar cane growers a means by 
which to identify it. 

Adult and larvae were examined using an 
incandescent light source (reflected light). 
Komerup and Wanscher (1978) was used as 
a color standard for the description of the 
adult. Genitalia were dissected as described 
by Clarke (1941), except Mercurochrome 
and chlorazol black were used as stains. 
Slide preparations were examined with dis- 
secting and compound microscopes. Mea- 
surements were made with a calibrated oc- 
ular micrometer. All specimens examined 
are deposited in The National Museum of 



VOLUME 101, NUMBER 1 



165 




1 



Fig. 1. Holotype of Blastohasis graminea. 



Natural History, Smithsonian Institution, 
Washington, D.C., (USNM), except where 
indicated otherwise. Label data taken ver- 
batim are expressed with quotations, while 
bracketed data are used to complete label 
data written in abbreviated form, or to help 
with the recognition of certain labels by de- 
scription of condition, e.g., [round label]. 

The ultrastructure of the larva was stud- 
ied with an Hitachi HH-S-2R scanning 
electron microscope at an accelerating volt- 
age of 20 kV. For SEM examination, larvae 
were fixed in 3% glutaraldehyde in 0.1 M 
potassium phosphate buffer (pH 7.3), rinsed 
in phosphate (pH 7.3), and postfixed in 2% 
osmium tetroxide in 0. 1 M potassium phos- 
phate (pH 7.3). After dehydration in ethyl 
alcohol, specimens were critical point dried, 
mounted on stubs with silver paint and 
paste, and coated with gold-palladium in a 
Polaron E5 1 00 sputter coater. 

Blastobasis graminea Adamski, 
new species 

(Figs. 1-26) 

Diagnosis. — Blastobasis graminea can 
be distinguished from other Blastobasis by 



the orange gray ground color, wide base of 
the uncus, rounded outer margin of the 
proximal flange, and wide ostial opening. 

Adult. — Head: Cephalic vestiture pale 
orange gray, except inner surface of labial 
palpus pale orange gray intermixed with 
brown scales tipped with white and few 
dark-brown scales, outer surface mostly 
brown intermixed with brown scales tipped 
with white, pale orange-gray scales, and 
dark-brown scales; segments paler near api- 
cal region. 

Thorax: Tegula and mesoscutum pale or- 
ange gray; legs with outer surface pale or- 
ange gray intermixed with orange-gray 
scales tipped with white, most specimens 
with foreleg and midleg with outer surface 
mostly grayish brown intermixed with 
grayish-brown scales tipped with white, or- 
ange-scales, and orange-gray scales tipped 
with white, leg segments and tarsomeres 
paler near apical region; forewing (Fig. 1), 
length 7.1-10.0 mm [n = 37], orange gray 
intermixed with orange gray scales tipped 
with white, brown scales tipped with white, 
and brown scales; several unrubbed speci- 
mens with discal cell region paler than out- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





Figs. 2, 3. Blastohasis graminea. 2, Wings, scale = 1.00 mm. 3. Male genitalia, scale = 0.5 mm. 



er region of wing; holotype with a brown midcell spot and two distal spots usually 

streak on basal part of posterior margin present; fringe scales mostly orange gray 

(Fig. 1); some specimens with veins de- tipped with white intermixed with orange 

marcated with white scales; a dark brown gray scales; undersurface grayish brown; 



VOLUME 101. NUMBER 1 



167 



cubitus four-branched, divergent from ra- 
dials and M, (Fig. 2); hindwing with both 
surfaces pale grayish brown; cubitus four- 
branched in a series typical of all New 
World Blastobasis (Fig. 2). 

Abdomen: Orange gray. 

Male Genitalia (Fig. 3): Uncus wide at 
base, posteriorly curved and narrowed api- 
cally, apical setae shorter than basal setae; 
gnathos bidentate; dorsal strut narrow; ter- 
gal setae numerous; diaphragma with mi- 
crotrichia throughout, extending to proxi- 
mal flange; proximal flange with stout mar- 
ginal setae, margin rounded; lower part of 
valva with marginal setae, numerous along 
apical third; juxta bandlike; aedoeagus api- 
cally rounded, with several stout anellar se- 
tae. 

Female Genitalia (Fig. 4): Ovipositor 
telescopic, in four membranous divisions; 
ostium within membranous area slightly 
posterior to seventh sternum; ostial opening 
wide; antrum membranous, narrowed 
abruptly anteriorly forming a common in- 
ception for ductus seminalis and ductus 
bursae; ductus bursae long, with two rows 
of platelike sclerotizations within anterior 
part; corpus bursae with posterior lobe near 
inception of ductus bursae; signum horn- 
like. 

Larva. — Length 6.5-14.9 mm [n = 207]. 
Body white, smooth, with head capsule, 
prothoracic shield, anal shield, pinacula and 
crochets yellowish orange. Head (Figs. 5- 
12, 17, 18): Hypognathous; epicranium 
smooth; adfrontal sclerites narrow, delim- 
iting frons dorsolaterally; frons closed; CI, 
C2, and C3 about equal in length, about 
three times length of Fl and F2; C3 closer 
to Fl than to C2; C2 slightly closer to mid- 
line than CI or C3; CI setae broadly 
curved, convergent; Fl subapical (Fig. 5); 
PI long, closer to P2 than A2; A2 closer to 
Al than A3; A3 nearly equidistant to A2 
and LI (Figs. 5-7); SI between stemmata 
2 and 3, and closer to S2 than to S3; SSI 
near mandibular articulation, and closer to 
SS2 than to SS3; SS2 between stemmata 5 
and 6 (Figs. 6, 7); labrum bilobed, each 




Fig. 4. Female genitalia of Blastobasis graminea. 
Scale = 1.00 mm. 



lobe with four subequal marginal setae and 
two subequal medial setae (Figs. 5, 6); 
mandibles slightly asymmetrical, with two 
distinct dentitions and two subequal setae 
on outer margin (Figs. 5, 6, 17); labium 
smooth with microtrichia along lateral mar- 
gin of proximal half; distal part of labium 
with median submental pit (Figs. 6, 8); la- 
bial palpus two-segmented, with dorsally 
directed subapical seta on basal segment. 
Sensilla types and arrangement on median 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



SSI "sS2 




Figs. 5-10. SEM of larva of Blastobasis graininea. 5, Frontal view of head capsule, 70x. 6, Ventral view 
of head capsule, VOX. 7, Lateral view of genal region of head capsule, 250 X. 8. Ventral view of labium. LBP 
= labial palpus, SPIN = spinneret, 250X. 9, 10, Sensilla on apex of maxillary palpus, A2 = sensillum stylo- 
conicum; Al, Ml, M2. LI, L2. L3, = sensilla basiconica; SD = sensilla digitiform, 2,500X. 



lobe and apex of palpus similar to that of 
Glyphidocera juniperella Adamski and 
Brown 1987, except for elongate depression 
near digitiform sensillum on part near L3 
sensillum (Figs. 8-10). Sensilla types on 



antenna (Fig. 1 1) similar to other Lepidop- 
tera (Schoonhoven and Dethier, 1966). Pro- 
thorax (Figs. 12, 18): Prothoracic shield 
with SDl and D2 about equal in length, 
twice length of XDl and XD2; SDl and D2 



VOLUME 101, NUMBER 1 



169 




Figs. 11-16. SEM of larva of Blastobasis graminea. 1 1, Sensilla on apical portion of antenna. 1 = sensilla 
basiconica; 2 = sensillum chaetica; 3 = sensullum styloconicum; 4 = sensillum trichodeum, 950X. 12. Lateral 
view of head capsule and thorax, 45X. 13, Ventral view of right proleg on A4, 200X. 14, A8-10, 60X. 15, 
Lateral view of AlO. SOX. 16, Posetrior view of AlO, SOX. 



about four times length of SD2 and Dl; Dl 
usually slightly longer than SD2; SD2 clos- 
er to SDl than to XD2; SD2 and D2 anter- 
iorad to Dl; LI about twice length of L2 
and L3; SVl about twice length of SV2; 



VI short (not illustrated). Mesothorax and 
metathorax (Figs. 12, 18): Dl anterodorsal 
to D2, on same pinaculum; D2 about three 
times length of Dl; SDl anterioventral to 
SD2, on same pinaculum; SDl about three 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

D2 



17 





SV1 




19 



20 




SD1 



Figs. 17-20. Lavva of Biastobasis gnimlnea. 17, Left mandible. 18, Lateral view of head capsule and thorax. 
19. A 1-3. 20, A7-10. 



times length of SD2; L2 anterioventral to 
LI, on same pinaculum; LI slightly longer 
than L2 and about three times length of L3; 
SVl about equal in length to LI and slight- 
ly caudal to L3; VI short (not illustrated). 
Abdomen (Figs. 13-16, 19, 20): Al and A2 
with D2 three times length of Dl; SDl 
about same length as D2; SD2 very short 
(Figured larger than normal), on same pin- 
aculum as SDl above spiracle; LI antero- 
dorsal to L2, L2 about twice length of LI; 
L3 caudal to D2, about equal in length to 
LI ; SV3 2-3 times length of SV2 and SVl ; 



SV2 and SVl in nearly straight line per- 
pendicular to longitudinal body axis, SV3 
slightly anterior to SVl; VI short (not il- 
lustrated); A3-A6 with SVl posterior to 
SV2; prolegs with crochets uniserial and 
triordinal, crochets smaller along outer mar- 
gin of planta; A7 with SV3 absent; A8 with 
SDl hairlike; one SV seta present; SVl 
nearly in verticle line with L3 and VI; SVl 
and L3 about equal in length, VI short (not 
illustrated); spiracle slightly larger than pro- 
thoracic and other abdominal spiracles; A9 
with SD2 absent; L3 ventral to L2; SVl 



VOLUME 101. NUMBER 1 



171 



caudal to D2; VI short (not illustrated); 
AlO (Figs. 14-16, 20): D2, SDl, and SD2 
about four times length of D 1 ; crochets uni- 
serial and triordinal. 

Holotype. — 9, "Colombia: Instituto Col- 
ombiano Agropecuario, Experiment Station 
"Palmira," Cauca Valley, 1 March-15 
March 1991, Ex. Sugar cane. Coll. Lucero 
Cardenas Duque, Emerged 21 April- 1 May 
1991." The holotype is not dissected and is 
deposited in USNM. 

Paratypes. — 3 ? , Same data as holotype. 
Paratypes are not dissected and are depos- 
ited in USNM. 

Other specimens examined. — COLOM- 
BIA: 2 c?, 7 9, "Miranda (Val.), en. cana 
a [zucar], Jul[y] 1984, L[ucero] Cardenas"; 
\ 6,1 9 from Vitor Becker Collection [yel- 
low label]; 1 6,5 9, "Miranda, VI-28- 
[19]84, L[ucero] Cardenas," "Tallos caiia 
de azucar," "$ Wing Slide by DA 3349, 
USNM 81585," [green label]," "9 Geni- 
talia Slide by D. Adamski 2885, USNM 
81422," [green label]; 3 9, "Riopaila, Par- 
asita Diatraea," "I-6-[19]65, 723-4," "II- 
1-[19]65, 9651-28," "II-15[19]65, 446- 
1"; 1 6, 1 9, "Ex. sugar cane, Ag. Exp. 
Sta., Palmira, Valle, Let. Oct. 3, 1941, B. 
Losada S," "9 Genitaha Slide by R. B. 
Selander, USNM 11, 157," [green label]; 2 
9, "Valle Ingenio del Cauca, H: caiia de 
azucar, Barrenador, Dic/[19]82, D-83," "9 
Genitalia Slide by D. Adamski 2849, 
USNM 81393," [green label], "9 Genitaha 
Slide by D. Adamski 2850, USNM 81394," 
[green label]; 2 6,1 9, "Valle Ing[enio] 
del Cauca, H: cana de azucar Col: L[ucero] 
Cardenas y Y.P Chacon, II-[19]83, D-83," 
"(? Genitalia Slide by D. Adamski 2847, 
USNM 81391," [green label], "d Genitalia 
Slide by D. Adamski 2846, USNM 81390," 
[green label], " 9 Genitalia Slide by D. 
Adamski 2848, USNM 81392," [green la- 
bel]. VENEZUELA: 2 9, "Tachira, El 
Cobre, 12,00 m[e]t[e]rs. May 1947," and 
"Tachira, La Grita, 1,450 m[e]t[e]rs, 
14. V. 1949," "Reared from larva in Sugar 
cane," "9 Genitaha Shde by R. B. Selan- 
der, USNM 11160," [green label], "9 Gen- 



italia Slide by R. B. Selander, USNM 
11161" [green label]; 2 6,1 9, "Maracay, 
450 m[e]t[e]rs, 28.iii.1949, H.E. Box", 
"February 1951," "June 1948," "Reared 
from larva in Sugar cane," "c? Genitalia 
Slide by D. Adamski 3038, USNM 81488" 
[green label], "d Wing Slide by J. G. 
Clarke, USNM 11209" [green label], "d 
Genitaha Slide by J. G. Clarke, USNM 
11209" [green label], "9 Genitalia Slide 
by R. B. Selander, USNM 11164" [green 
label]; 1 9, "Carabobo, Cent. Tacarigua, 
450 m[e]t[e]rs, September 1947, H.E. 
Box," "Reared from larva in Sugar cane," 
"9 Genitalia Slide by R. B. Selander, 
USNM 11159" [green label], 1 6,3 9, 
"Yaracuy, Chivacoa, 230 m[e]t[e]rs, Feb- 
ruary 1950," "28. ii. 1950," "San 
Pa[illegible], 400 m[e]t[e]rs, 22. ii. 1949, 
H.E. Box," "Reared from larva in Sugar 
cane," "d Genitalia Slide by D. Adamski 
3037, USNM 81487," [green label], "9 
Genitalia Slide by R. B. Selander, USNM 
11163" [green label], "Reared from larva 
in Sugar cane," " 9 Genitalia Slide by J. G. 
Clarke, USNM 11210" [green label], 
"Reared from larva in Sugar cane," "9 
Genitalia Slide by R. B. Selander, USNM 
11162," [green label], "Reared from larva 
in Sugar cane"; 1 6, "Merida, nr. Egido, 
1,500 m[e]t[e]rs, 8.VI.1949, H. E. Box," 
"d Genitaha Slide by D. Adamski 3039, 
USNM 81489" [green label]; 1 9, "Miran- 
da, Sta. Lucia, 180 m[e]t[e]rs, 5.iii.l948, 
H.E. Box," "9 Genitaha Slide by J. C. 
Clarke, USNM 11207" [green label], 
"Reared from larva in Sugar cane"; 1 9, 
"Aragua, El Conseja, 550 m[e]t[e]rs, 
March 1951, H.E. Box," "Reared from lar- 
va in Sugar cane," "9 Genitalia Slide by 
R. B. Selander, USNM 11158" [green la- 
bel]; 1 6, "El Limon, nr, Maracay, 460 
m[e]t[e]rs, 31.iii.l950, H.E. Box," "Reared 
from larva in Ciox lochryma-jobi [L]"; 1 
9, "Zulia, Perijo, Mts. close to Colombia, 
Dec. 1950, E Fernandez Yepoz," "Reared 
from larva in Setaria paniculifera [Four- 
nier]," " 9 Genitalia Shde by J. F. Clarke, 
USNM 11208" [green label]; 1 9, "Vene- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



zuela, Turbio Valley, nr. Barquisimeto, 
1956, P. Guagliumi, Larva boring sugar- 
cane, COM. INST. ENT. COLL. NO. 
1521 1, Press[ented] by Com. Inst. Ent., BM 
1957-256." Fourteen additional adult spec- 
imens were examined at The Natural His- 
tory Museum, London, with above label 
data. Larvae studied were collected and 
preserved in alcohol with the following 
data, "Colombia: Instituto Colombiano 
Agropecuario, Experiment Station "Palmi- 
ra," Cauca Valley, 15 December-20 Janu- 
ary 1990, Ex. Sugar cane. Coll. Lucero Car- 
denas Duque." All larval specimens are de- 
posited in the USNM alcohol collection. 

Types examined. — Lectotype designated 
by Clarke, 6, Blastobasis obstricta Mey- 
rick, "Lectotype" [round label], "Bartica, 
Brit[ish] Guiana, Parish 1.13," "Lectotype, 
Auximobasis obstricta Meyrick, J.F.C.C. 
1948," "(? Genitalia on Slide 5-X-1948, 
J.F.G.C. 8078," Auximobasis obstricta 
Meyr., E. Meyrick det., in Meyrick Coll. 
21/1," ""obstricta Meyr.," "Meyrick Coll., 
BM 1938-290," [Natural History Museum, 
London, England]. Lectotype, S, Blasto- 
basis subolivacea Walsingham, "S[aint] 
Thomas, 9.IV.[18]94" [hand-written pink 
label], ""Blastobasis subolivacea 125.2089 
WLSM, S, TYPE" [hand-written label], 
""S Genitalia Slide by D. Adamski, 3470" 
[green label], "Holotype, Blastobasis su- 
bolivacea Wlsm, d," ["Grigore Antipa" 
National Museum of Natural History, Bu- 
charest, Romania]. 

Etomology. — Blastobasis graminea is 
named after the plant family Gramineae be- 
cause larvae feed on several grass hosts. 

Discussion 

Auximobasis obstricta Meyrick 1918, is 
transferred to Blastobasis Zeller, 1855, and 
Blastobasis subolivacea Walsingham, 1897, 
is transferred to Holcocera Clemens 1863 
(new combinations). 

Blastobasis graminea is probably more 
closely related to Blastobasis obstricta 
Meyrick, 1918, n. comb, than to any other 
described Blastobasis. Both species differ 



markedly in wing pattern and in several 
male and female genitalic features. How- 
ever, males of both species share an uncus 
with a widened base, a bidentate gnathos, 
and stout marginal setae along the outer 
margin of the proximal flange. Females 
share a widened ostium. 

Martorell (1976) reported two species of 
Blastobasidae feeding within sorghum 
heads in the Vieques Islands east of Puerto 
Rico, but I have not seen any specimens to 
substantiate this. 

Biology 

Cardenas and Hernandez (1985) de- 
scribed the biology of Blastobasis grami- 
nea on sugar cane in Colombia; these find- 
ings are summarized below. The most se- 
vere damage by B. graminea occurs within 
the terminal third of the sugar cane plant, 
however, damage can occur in lower re- 
gions as well. When the damage is extreme 
the apical portion of the plant dies. 

Early instars of B. graminea feed on the 
surface tissue layers. When the larva is 
able, it bores into the stem. Galleries are 
usually irregularly shaped (Figs. 22-24); 
the larvae never excavate more than two 
internodes of the plant. Mature larvae usu- 
ally pupate between the stem and the sheath 
(Fig. 21), but sometimes within the stem 
(Fig. 22). In addition to sugar cane, larvae 
feed on corn (Figs. 25, 26), sorghum, Coix 
lacryma-jobi L., and Setaria paniculifera 
Foumier. 

There appears to be a strong correlation 
(Ratio of 8:1) between the presence of Dia- 
traea saccharalis and Blastobasis grami- 
nea, however, it is not known which moth 
species attacks the plant first. 

Acknowledgments 

I thank Ingeborg Zenner-Polania, former 
director, Programa de Entomologia, Insti- 
tuto Colombiano Agropecuario, Bogota, 
Colombia, for the coordination of activities 
related to the acquisition of specimens of 
Blastobasis graminea: Lucero Cardenas 
Duque, of the above institution, for live and 



VOLUME 101. NUMBER 1 



173 




Figs. 21-26. Larval damage of Blastohasis grominea. 21, Pupal site on sugar cane (see arrow). 22, Pupa 
within sheath of sugar cane (see anow). 23, Larvae and damage within sugar cane stem (see arrow). 24, Larval 
gallery within sugar cane stem (see arrow). 25, Larval damage in corn stem (see anow). 26, Larva within corn 
stem (see arrow). 



preserved larvae, and photographic prints 
of larval damage; Klaus Sattler, Michael 
Shaffer and Kevin Tuck, of the Natural 
History Museum, London, England, for 
their help with examination and photog- 
raphy of type specimens; Dorel Rusti, 
"Grigore Antipa" National Museum of 
Natural History, Bucharest, Romania, for 
the loan of the lectotype of Blastohasis su- 
bolivacea Walsingham; Greta Tyson and 
Michael Sullivan, of the Electron Micro- 



scope Center, Mississippi State University, 
for their help with the preparation of the 
specimens and photographic plates; Carl 
Hansen of the Office of Imaging, Printing 
and Photographic Services for the photo- 
graph of the holotype; the late John E 
Gates Clarke, Smithsonian Institution, for 
referring this research problem to me. This 
research was supported in part by NSE 
Grant BSR85-01212 and a grant from Sig- 
ma Xi. 



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Literature Cited 

Adamski. D. and R. L. Brown. 1987. A New Nearctic 
Glyphidflcera with descriptions of all stages (Lep- 
idoptera: Blastobasidae: Symmocinae). Proceed- 
ings of the Entomological Society of Washington 
89(2): 329-343. 

. 1989. Morphology and Systematics of North 

American Blastobasidae (Lepidoptera: Gelechioi- 
dea). Mississippi Agricultural Forest Experiment 
Station Technical Bulletin 165. Mississippi Ento- 
mological Museum Publication No. 1. 70 pp. 

Box, H. E. 1953. List of Sugar-Cane Insects: A Syn- 
onymic Catalogue of the Sugar-cane Insects and 
Mites of the World, and of their Insect Parasites 
and Predators, Arranged Systematically. Com- 
monwealth Institute of Entomology, London, 100 
pp. 

Cardenas Duque, L. and Miria del Pilar Hernandez. 
1985. Barrenador de la caria de azucar en Colom- 
bia. Miscelanea, Sociedad Colombiana de Ento- 
mologia 1: 12-17. 

Clarke, J. EG. 1941. The preparation of slides of the 
genitalia of Lepidoptera. Bulletin of the Brooklyn 
Entomological Society 36: 149-161. 

Clemens, B. 1863. American micro-lepidoptera. Pro- 
ceedings of the Entomological Society of Phila- 
delphia 2: 119-129. 

Guagliumi, P. 1962. Las plagas de cana de azucar en 



Venezuela. Ministerio de Agricultura y Cria. Ma- 
racay, Venezuela. Monografi'a no. 2. 2 partes. 789 
pp. 

Hodges, R. W. In press. Gelechioidea. //; Kristensen, 
N. P., ed., Handbuch der Zoologie. 

Kornerup, A. and J. H. Wanschner. 1978. Methuen 
Handbook of Colour. Third Ed. Methuen and Co., 
Ltd.. London. 252 pp. 

Martorell. L. F. 1976. Annotated Food Plant Catalogue 
of the Insects of Puerto Rico. Agricultural Exper- 
iment Station, University of Puerto Rico, Depart- 
ment of Entomology. 303 pp. 

Meyrick, E. 1894. On a collection of Lepidoptera from 
upper Burma. Transactions of the Entomological 
Society of London 1894: 1-29. 

. 1918. Exotic Microlepidoptera. 2. pp. 385- 

480 (1916-23) Taylor and Francis, London. Re- 
printed by C. W. Classey. Ltd.. 1969. 640 pp. 

Schoonhoven L. M. and V. G. Dethier. 1966. Sensory 
aspects of host-plant discrimination of lepidopter- 
ous larvae. Archives Neerlandaises de Zoologie 
16(4): 497-530. 

Walsingham, Lord (Thomas de Grey). 1897. Revision 
of the West Indian micro-Lepidoptera with de- 
scriptions of new species. Proceedings of the Zoo- 
logical Society of London 1897: 54-183. 

Zeller, R C. 1855. Die Arten der Gattung Butalis be- 
schreiben. Linnaea Entomologica 10: [169]-269. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 175-196 

A REVISION OF THE FLAVIDUS GROUP OF THE 
GENUS CHRYSOPS MEIGEN (DIPTERA: TABANIDAE) 

Tevis M. Baier 

Department of Zoology, University of New Hampshire, Durham, NH 03824, U.S.A. 
(e-mail: tbaier@jlc.net) 



Abstract. — A taxonomic revision of the Chrysops flavidus species group is provided 
with a key to species, descriptions, distribution maps, and illustrations of wings and an- 
tennae for each of the nine species currently recognized in this group. The relationships 
and identification of these nine species are based on analysis of morphological similarity 
involving 13 critical characters. A total of 4,843 specimens from 14 institutions were 
examined during this study. Two new species in the Chrysops flavidus group are described, 
Chrysops sandyi and Chrysops tumidicornis, both previously misidentified as Chrysops 
atlanticus. The male of Chrysops dixiauus is described for the first time. 

Key Words: Diptera, Tabanidae, Chrysops flavidus group, key to species 



The genus Chrysops Meigen is currently 
represented by 87 described species in the 
Nearctic Region north of Mexico (Burger 
1995), and 291 world wide. This genus was 
last revised in North America by Philip 
(1955). Philip's revision included a key to 
95 species and 14 subspecies, with corre- 
sponding notes on these taxa, and descrip- 
tions of 4 new species and 5 new subspe- 
cies (but it also treated species from Central 
America and the Caribbean). 

Although no one has divided all of the 
Nearctic Chrysops into definitive species 
groups, some species groups have been rec- 
ognized by tabanid taxonomists. The three 
commonly used are the Chrysops callidus 
group, the C. carbonarius group, and the C. 
flavidus group (Pechuman 1949). The C. 
flavidus group is particularly difficult be- 
cause of the large amount of intraspecific 
variation within the included taxa. Al- 
though three species have been described in 
this group during the past 45 years, no one 
has studied the group as a whole. 

The Chrysops flavidus group currently 



includes 7 previoously described species 
and two new species: Chrysops atlanticus 
Pechuman, C brunneus Hine, C celatus 
Pechuman, C dixiauus Pechuman, C flav- 
idus Wiedemann, C pudicus Osten Sacken, 
and C. reicherti Fairchild, C. sandyi, n.sp., 
and C. tumidicornis, n. sp. I will redescribe, 
clarify relationships and summarize the bi- 
ology and geographic distribution of this 
group. Immature stages and habitat are dis- 
cussed under the description of each spe- 
cies. 

Materials and Methods 

I examined 4,843 specimens, (4,684 fe- 
males and 159 males), from the collections 
listed below. The acronyms are those of Ar- 
nett et al. (1993), except for the University 
of New Hampshire Collection (UNHC). 

AMNH: Department of Entomology, 
American Museum of Natural 
History, New York, NY; David A. 
Grimaldi. 

BMNH: Department of Entomology, The 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Natural History Museum, Lon- 
don, U. K.; John E. Chainey. 

CASC: Department of Entomology, Cali- 
fornia Academy of Sciences, San 
Francisco; Paul H. Arnaud, Jr. 

CNCI: Canadian National Collection, 
Centre for Land and Biological 
Resources Research, Agriculture 
Canada, Ottawa, Ontario; J. M. 
Cumming. 

CUCC: Department of Entomology, 
Clemson University, Clemson, 
SC; Michael A. Floyd. 

CUIC: Department of Entomology, Cor- 
nell University, Ithaca, NY; E. Ri- 
chard Hoebeke. 

FMNH: Division of Insects, Field Museum 
of Natural History, Chicago, IL; 
Alfred F. Newton, Jr. 

FSCA: Florida State Collection of Ar- 
thropods, Division of Plant Indus- 
try, Gainesville; G. B. Fairchild. 

INHS: Illinois Natural History Survey 
Insect Collection, Champaign; 
Kathleen R. Methven. 

OSUC: Ohio State University, Collection 
of Insects and Spiders, Columbus; 
Andrey Sharkov. 

TAMU: Department of Entomology Insect 
Collection, Texas A. & M. Uni- 
versity, College Station; Edward 
G. Riley. 

UGCA: Entomology Collection, Universi- 
ty of Georgia, Athens; Cecil L. 
Smith. 

UNHC: Entomological Museum, Depart- 
ment of Zoology, University of 
New Hampshire, Durham; Donald 
S. Chandler. 

USNM: National Museum of Natural His- 
tory, Smithsonian Institution. 
Washington, DC; R. V. Peterson. 

The following insect collections are also 
cited in this paper: 

MNHN: National Collection of Insects, 
Museum National d'Histoire Na- 
turelle, 45, Rue Buffon, Paris 
75005 France. 



MRSN: Museo Regionale Scienze Natur- 
aH, Via Gioletti 36, Torino 10128, 
Italy. 

MZLU: Museum of Zoology, Lund Uni- 
versity, Helgonavagen 3, S-223 
62 Lund, Sweden. 

ZMHB: Museum fiir Naturkunde der 
Humboldt Universitat zu Berlin, 
Bereich Zoologischer Museum, 
InvalidenstraBe 43, 1040 Berlin, 
Germany. 

The Chrysops flavidus group has few dis- 
tinctive structural characters, and the spe- 
cies can be difficult to identify. Color pat- 
terns are important in identification but 
must be used with caution because of var- 
iation. The chaiacters described below are 
the most useful in separating the species. 
The morphological terminology used in this 
paper follows that used by Teskey (1990). 
Principal characteristics used are: body 
length, in millimeters, from the base of the 
antennae to the apex of the abdomen; the 
size and shape of the scape and pedicel as 
well as the length ratio and color of the 
scape, pedicel, and flagellum; shape of the 
frontal callus; color of the mesoscutum; 
wing pattern, particularly the width of the 
apical spot, the location of the outer margin 
of the crossband, and the infuscation of the 
5th posterior cell; color of the hind femur; 
and abdominal color pattern. 

For illustrations, I chose a specimen that 
most closely resembled the description of 
the holotype. One wing from each specimen 
was removed, placed between two 2X2 
glass slides and scanned into a Dell 486p/ 
25 computer using MICROTEK Scan- 
Maker 35t. The scanned picture was printed 
and used as a template for the final drawing. 
The antennal drawings were produced with 
a camera lucida. 

The locality data from all specimens ex- 
amined were entered into a database using 
Wordtech Systems, dBXL®, (a dBASE III 
Plus® compatible format). The information 
collected was placed into 9 "fields": spe- 
cies, country, state, county, town, collector. 



VOLUME 101. NUMBER 1 



177 




apical spot 



hyaline triangle 



flagellum pedicel 



scape 




crossband 
5th posterior cell 



basal callus 



antennal socket 



ocelli 




Figs. 1-3. 1. Wing of Chrysops species. 2, Antenna of Chiysops species. 3, Anterior view of head of 
Chrysops species. 



museum, count (number of specimens with 
identical data), and sex (male or female). 
Locality data for specimens I did not ex- 
amine were taken from the distribution re- 
cords of L. L. Pechuman, now stored in the 
Zoology Department at the University of 
New Hampshire. Distribution maps were 
then produced for each species from the 
above database and from the records of L. 
L. Pechuman. 

Diagnosis of the 
Chrysops flavidus Group 

Frontoclypeus glossy yellow with no 
mid-facial stripe; frontal callus yellow to 
light brown, occasionally with brown upper 
margin; frons yellow pollinose; mesoscu- 
tum with 3 longitudinal stripes, darker than 
ground color; proepimeron and proepister- 
nal callosity yellow tomentose with long 
yellow hair; wing pattern with broad apical 
spot entering 2nd submarginal cell, cross- 
band reaching hind margin of wing, hyaline 



triangle not crossing vein R2+3, cell br at 
least V3 infuscated basally; abdominal ter- 
gites 2-4 patterned with median inverted 
"V" set over pale median triangle. 

Key to Species of the 
Chrysops flavidus Group 

1. Scape and pedicel distinctly swollen, together 
longer than flagellum (Fig. 14). Frontal callus 
twice as wide as high, with no black markings. 
Abdomen with inconspicuous median trian- 
gles. Wing pattern with extensive apical spot, 
reaching crossband posteriorly and enclosing 
hyaline crescent (Fig. 23) brunneus Hine 

- Scape and pedicel less swollen, combined 
length equal to or shorter than flagellum. With- 
out remaining combination of characters .... 2 

2. Usual hyaline areas of wing slightly smoky to 
very smoky, outer margin of crossband angled 
from vein R4 to middle of vein M4 (Fig. 22). 
Scape and pedicel moderately swollen, com- 
bined length equal to or rarely longer than fla- 
gellum (Fig. 13). Mesoscutum green-gray iri- 
descent with black longitudinal stripes . . . 

atlanticiis Pechuman 

- Without above combination of characters ... 3 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



3. Mesoscutum greenish gray or steel gray in 
ground color, longitudinal stripes black. Outer 
margin of crossband sinuous 4 

- Mesoscutum yellow in ground color, longitu- 
dinal stripes brown. Outer margin of cross- 
band, straight, concave, or sinuous 6 

4. Antenna not swollen, flagellum at least Vs lon- 
ger than scape and pedicel combined (Fig. 15). 
Frontal callus spade shaped, width and height 
subequal. Without dark spot under scutellum 

celatus Pechuman 

- Scape and pedicel slightly swollen to swollen, 
combined length subequal to flagellum. Frontal 
callus, width and height not equal. Abdomen 
with dark spot under scutellum, often extend- 
ing to lateral edges of tergite 1 5 

5. Hyaline triangle not reaching vein R^^,, cells 
br and bm % and V3 infuscated basally (Fig. 30). 
Scape and pedicel swollen, pedicel barrel 
shaped (Fig. 21). Frontal callus light brown. 
Mesoscutum gray green with black stripes . . 

tumidicomis. n. sp. 

- Hyaline triangle reaches vein R2 + 3, apical spot 
narrowly enters 2nd submarginal cell, cells br 
and bm Vi and '/^ infuscated basally (Fig. 27). 
Scape and pedicel slightly swollen (Fig. 18). 
Frontal callus reddish brown with black upper 
margin. Mesoscutum steel gray with black 
stripes pudicus Osten Sacken 

6. Abdominal tergites 1-2 light yellow to straw 
color, dark inverted "V" on tergite 2 com- 
pressed, not reaching beyond the middle of ter- 
gite 2. Flagellum 1.3 to 1.5 times longer than 
scape and pedicel combined 7 

- Abdominal tergites 1-2 yellow, orange or light 
brown, inverted "V" pattern not compressed, 
extending beyond the middle of tergite 2 to- 
ward anterior margin. Flagellum equal to or up 
to 1.2 times longer than scape and pedicel 
combined 8 

7. Fifth posterior cell infuscated, outer margin of 
crossband sinuous, cells br and bm V3 and Va 
infuscated basally (Fig. 28). Hind femur red- 
dish-brown. Tergites 1-2 straw-colored, medi- 
an anterior portion of tergite 2 with a greenish 
cast. Length 8-9.5 mm reicherti Fairchild 

- Fifth posterior cell hyaline, vein Cu, infuscated 
at border, outer margin of crossband straight or 
convex, cells br and bm Vi and '/, infuscated 
basally (Fig. 25). Hind femur dark brown. Ter- 
gites 1-2 yellow without a greenish cast. 
Length 6-8.5 mm dixianus Pechuman 

8. Hind femur yellow. Mesoscutum yellow with 
brown stripes. Outer margin of crossband 
straight or concave, cells br and bm V2 and V3 
infuscated basally (Fig. 26). Scape and pedicel 
slightly swollen (Fig. 17). Length 8-10 mm 

fiavidus Wiedemann 



- Hind femur dark brown basally. Mesoscutum 
yellow green with brown stripes. Outer margin 
of crossband straight, hyaline triangle slightly 
tinted, cells br and bm V3-V2 and 'A infuscated 
basally (Fig. 29). Scape and pedicel not swol- 
len (Fig. 20). Length 5.7-7.5 mm 

sandyi, n. sp. 

Chrysops atlanticus Pechuman 
(Fig. 4, 13, 22) 

Chrysops canifrons Walker 1848: 197-198. 
Holotype female: Florida (BMNH). Se- 
nior synonym of atlanticus, suppressed 
by the International Commission on Zoo- 
logical Nomenclature, Opinion No. 1711, 
1993. 

Chrysops atlanticus Pechuman 1949: 79- 
82. Holotype female: Rehoboth Beach, 
Delaware (CUIC); Jamback and Wall 
1959: 23-24, description of egg, larva 
and pupa; Teskey 1969: 29-30, descrip- 
tion of larva; Goodwin 1972: 104, de- 
scription of pupa. 

Diagnosis. — Length 6.5-9.2 mm. Scape 
and pedicel moderately swollen, flagellum 
equal to length of scape and pedicel com- 
bined. Mesoscutum grayish green in ground 
color. Wing pattern with smoky tinge, hy- 
aline areas not clearly defined, hyaline tri- 
angle not extending beyond lower half of 
1st submarginal cell. 

Female. — Light to dark brown, length 
8.5-9.2 mm. Scape and pedicel moderately 
swollen, light brown, basal flagellomere 
light brown, apical flagellomeres black, 
length of flagellum subequal to combined 
length of scape and pedicel, antennal ratio 
15:10:25. Frontoclypeus glossy yellow, eye 
margins bearing yellow tomentum that ex- 
tends medially beneath tentorial pits to 
frontoclypeal suture. Maxillary palp brown 
with sparse black hairs. Frons yellow and 
gray tomentose with yellow hairs at vertex, 
width 1.13 times height. Frontal callus ob- 
long, bulbous, light brown, width 1.54 
times height. Vertex with glossy black in- 
tegument surrounding shining brown ocelli. 
Mesoscutum grey green iridescent with 
three black longitudinal stripes, sublaterals 



VOLUME 101, NUMBER 1 



179 




Figs. 4-9. Distributions. 4, Chtysops atlanticus. 5, C. brunneiis. 6, C. celafus. 7. C. dixianus. 8. C. fiavidiis. 
9, C pudicus. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



wider than median one, scutellum predom- 
inantly brown, lighter medianly. Legs pre- 
dominantly yellow, hind coxa and fore ta- 
rusi brown, fore tibia with black hair, mid 
and hind tibia with mixed yellow and black 
hair. Wing pattern with usual hyaline areas 
smoky, apical spot broad, fading into 2nd 
submarginal cell, crossband brown with ir- 
regular outer margin, crossing 1st posterior 
cell at its center, a perpendicular line drawn 
from vein R4 to hind margin of wing not 
forming line parallel to outer margin of 
crossband, hyaline triangle not extending 
beyond posterior Vi of 1st submarginal cell, 
discal cell, 4th and 5th posterior cells in- 
fuscated, 5th posterior cell paler in center, 
cells br and bm % and V3 basally infuscated. 
Halter brown stalk yellow-brown knob. 
First abdominal tergite dark yellow, black 
haired medianly, tergites 2-5 with subme- 
dian oblique dark spots, gray posterior mar- 
gin extending medianly into pale orange, 
equilateral triangle, dark spots on tergite 2 
outline triangle and join anteriorly forming 
inverted "V" pattern, tergites 3-5 with sub- 
median dark spots slightly separated ante- 
riorly. Sternites 1-2 yellow, sternite 3 mot- 
tled yellow and brown, sternites 4-7 dark 
brown. 

Male. — Similar to female except for usu- 
al sexual differences and following char- 
acteristics. Length 6.5-8 mm. Scape and 
pedicel slightly more enlarged with black 
hair that is longer and finer than on female. 
Mesoscutum sparsely covered with long 
yellow hair. Cell br entirely infuscated ex- 
cept for a subapical hyaline spot. Abdomen 
with yellow and black hair scattered over 
pattern. 

Material examined. — 934 9 and 18 6 
examined from the following collections: 
CASC, CNCL CUCC, CUIC, FMNH, 
FSCA, INHS, OSUC, TAMU, UGCA, 
UNHC, USNM. 

Distribution. — Atlantic coast of North 
America from Maine to Florida. I have ex- 
amined specimens from AL, DE, FL, GA, 
LA, MA, ME, MD, MS, NC, NH, NJ, NY, 



SC, TN, and VA. There are also published 
records for CT and RL 

Biology. — The larvae have been found in 
salt marsh and brackish pools. Based on 
their narrow distribution along the coast, 
they are probably restricted to this saline 
environment. Flight times for C. atlanticus 
are between April and September in Loui- 
siana (Tidwell 1973), June to September in 
Virginia (Pechuman 1973), and late May to 
mid October in New York (Pechuman 
1981). The adults are abundant during June 
and July throughout their range and are eco- 
nomically important due to their aggressive 
biting behavior. Thirty C atlanticus bites 
were counted on 1 human in 90 seconds in 
a cultivated field near Cedarville, New Jer- 
sey, (Hansens, 1980). Anderson (1971) de- 
scribed C. atlanticus as being autogenous, 
depositing the first egg mass before seeking 
a blood meal. Subsequent work by Mag- 
narelli and Anderson (1976) concluded that 
populations of C atlanticus can be main- 
tained by the first oviposition, making them 
difficult to control. 

Chrysops atlanticus is not known to 
transmit any disease agent in nature; how- 
ever, in the laboratory, tests have demon- 
strated it to be an effective experimental 
vector of the African filarial worm Loa loa 
(Orihel and Lowrie 1975). 

Discussion. — Chrysops atlanticus is 
most similar to C. sandyi, C. tumidicornis, 
and to C. brunneus, based on the smoky 
wing pattern, association with a saline en- 
vironment, and swollen antennae. Chrysops 
atlanticus differs from C brunneus in its 
narrower scape and pedicel (Fig. 22) that 
when combined are equal in length to the 
flagellum, moderately inflated frontal cal- 
losity, grayish-green mesoscutum, and wing 
pattern without clearly defined hyaline ar- 
eas. Chrysops atlanticus differs from C 
sandyi in its larger size, blackish longitu- 
dinal stripes on the mesoscutum, and cross- 
band with irregular outer margin. Chrysops 
atlanticus differs from C tumidicornis in its 
larger size, less swollen pedicel, smokier 
wing pattern, and lighter abdominal pattern 



VOLUME 101, NUMBER 1 



181 



that does not have a dark spot on tergite 1 
under the scutellum. There appears to be no 
overlap in the geographical distribution of 
C. atlanticiis with either C. sandy! or C. 
tumidicornis. 

Color variation in C. adanticus is com- 
mon throughout its range, making it partic- 
ularly difficult to define. The abdominal 
pattern can vary from tergites 2-6 having a 
pale inverted "V" formed by hair, to a 
dark, well defined, integumental inverted 
"V" pattern that includes Vs or more of the 
tergite. In all these cases, the characteristic 
wing pattern described above and the in- 
flated scape and pedicel will define C. at- 
lanticus. 

Chrysops brunneus Hine 
(Figs. 5, 14, 23) 

Chrysops brunneus Hine 1903: 34. Syn- 
types female and male: Sandusky, Ohio, 
9 S (OSUC), 1 $ (BMNH); Goodwin 
1976: 343, description of larva and pupa. 

Diagnosis. — Robust species, length 8-10 
mm. Antenna with scape and pedicel swol- 
len, flagellum distinctly shorter than com- 
bined length of scape and pedicel. Frontal 
callus strongly inflated, width twice height. 
Wing with apical spot extensive, reaching 
around margin of wing to crossband and 
enclosing hyaline crescent. Cells br and bm 
% and Vi infuscated basally. Abdominal pat- 
tern indistinct. 

Female. — Yellowish brown. Scape and 
pedicel distinctly swollen, scape twice as 
long as broad, light brown, black setose, 
basal flagellomere yellow basally, dark 
brown apically, apical flagellomeres black, 
flagellum distinctly shorter than scape and 
pedicel combined, antennal ratio 18:12:25. 
Frontoclypeus glossy yellow, eye margins 
bearing yellow tomentum that extends me- 
dianly beneath tentorial pits to frontocly- 
peal suture. Maxillary palp light brown 
with sparse black hairs. Frons yellow to- 
mentose with scattered yellow hairs, con- 
vergent above, height 1.3 times width at 
base. Frontal callus elliptical, distinctly nar- 



rowed laterally, strongly inflated, light 
brown, upper margin dark brown, width 
twice height. Vertex dark brown tomentose, 
except glossy around each ocellus, ocelli 
dark brown. Mesoscutum light brown to- 
mentose and yellow pilose, except lateral 
margins pale yellow-gray tomentose, bear- 
ing 3 longitudinal stripes, median stripe 
very narrow, dark brown, sublateral stripes 
brown and broad. Scutellum dark brown 
basally, brown apically, yellow pilose. 
Pleuron yellow-gray tomentose. Fore coxa 
light brown tomentose, mid and hind coxa 
dark brown tomentose, femur, tibia and tar- 
sus light brown, femur and tibia bearing 
yellow hairs, except apices of tibia bearing 
mixed yellow and black hairs, tarsus bear- 
ing black hairs. Wing pattern distinct, apical 
spot broad, extending around wing margin 
to crossband, leaving narrow hyaline band 
along outer margin of crossband that does 
not reach vein R2f3, outer margin of cross- 
band straight or sinuous, 5th posterior cell 
infuscated, cells br and bm % and Vi their 
length respectively. Halter with dark brown 
stalk and light brown knob. Abdomen uni- 
formly brown, lacking dark integumental 
markings, tergite 1 with black hair medi- 
anly, yellow hair laterally, tergites 2-4 pre- 
dominantly black-haired, except for yellow 
hairs on posterior margin that expand me- 
dianly into indistinct yellow-haired trian- 
gles, tergites 5-7 with mixed black and yel- 
low hairs forming no distinct pattern. Ster- 
nites 1-5 light brown, sternites 3-5 with 
progressively larger median and lateral 
brown integumental spots, 6-7 dark brown, 
all sternites with mixed black and yellow 
hairs. 

Male. — Resembles female except for the 
usual sexual differences and the following 
characteristics. Length 8.2-10 mm, scape 
and pedicel slightly more inflated than in 
female, with longer, finer black hair, fron- 
toclypeus with yellow tomentose triangle at 
vertex, cheeks yellow pollinose. Long yel- 
low hairs scattered over mesoscutum, scu- 
tellum, and abdomen. 

Material examined. — 502 9 and 14 S 



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11 



J"M:\ 



Figs. 10-12. Distributions. 10, Chiysops reiclierti. 11, C. sandyi. 12, C. tiiinidicumis. 



were examined from the following collec- 
tions: AMNH, CASC, CNCI, CUCC, 
CUIC, FMNH, FSCA, INHS, OSUC, 
TAMU, UGCA, UNHC, USNM. 

Distribution. — New York to Florida, 
west to Texas, and north to Michigan and 
southern Ontario. It is not found in the Ap- 
palachian Highlands and surrounding ter- 
rain. I have examined specimens from: AL, 
AR, DE, FL, GA, IL, IN, KS, KY, LA, MD, 
MI, MS, NJ, NY, OH, OK, PA, SC, TN, 
TX, and VA. It is also known from lA, MO, 
NC, and Ontario, Canada. 

Biology. — Chrysops brunneus is associ- 
ated with both fresh and salt water habitats. 
Goodwin (1976) reared a larva of this spe- 
cies collected from a salt marsh in Mc- 
Clellanville, SC. Adults are commonly 



found in marshes near lakes such as Lake 
Michigan, Lake Erie and Lake Ontario. 

Discussion. — Chrysops brunneus is very 
distinctive and can be separated from other 
species of the C flavidus group by its 
strongly inflated scape and pedicel that 
combined are longer than the flagellum, and 
by its reduced hyaline triangle appearing as 
c' crescent-shaped area between the cross- 
band and broad apical spot. The smoky 
wing pattern of C. brunneus and its pres- 
ence in a saline environment are similar to 
that of C. atlanticus, but is easily separated 
by those characters noted above. 

The only variation observed in this spe- 
cies was the abdominal pattern that has ter- 
gites 2-3 with small, median, black dashes 
on the integument in approximately 5% of 



VOLUME 101, NUMBER 1 



183 




atlanticus 

13 



brunneus 

14 



ce/atus 

15 




dixianus 



16 



reicherti 



19 



flavidus 

17 




sandyi 

20 



pudicus 



18 




tumidicornis 

21 



Figs. 13-21. Female antennae. 13, Clvysops atlanticus. 14, C hnmneus. 15, C. celatus. 16, C. dixianus. 
17, C. flavidus. 18, C. pudicus. 19, C reicherti. 20, C sandyi. 21, C tumidicornis. 



the specimens examined. Ordinarily the ab- 
dominal pattern is indistinct with only some 
dark shadowing from dark hairs. 

Chrysops celatus Pechuman 
(Figs. 6, 15, 24) 

Chrysops flavidus celatus Pechuman 1949: 
82-83. Holotype female: Medford Lakes, 
New Jersey (CUIC); Pechuman 1957: 30, 
description of male. 

Chrysops celatus: Teskey 1969: 34, de- 
scription of larva and pupa. 

Diagnosis. — Black and yellow, length 8- 
9.8 mm. Scape and pedicel not swollen, fla- 
gellum V3 longer than scape and pedicel 
combined. Frontal callus spade shaped, 
width and height subequal. Mesoscutum 
greenish gray iridescent with black longi- 
tudinal stripes. Cells br and bm Vi and '/, 
infuscated basally. 

Female. — Scape and pedicel not swollen, 
light brown, black setose, basal flagellomere 
light brown basally, black apically, apical fla- 
gellomeres black, 1.3 times length of basal 
flagellomere, antennal ratio 15:10:34. Fron- 
toclypeus glossy yellow, eye margins bearing 
yellow tomentum that extends medianly be- 



neath tentorial pits to frontoclypeal suture. 
Maxillary palp and labrum yellow. Frons pre- 
dominantly yellow-gray tomentose with scat- 
tered yellow hair, slightly convergent above, 
height 1 .5 times width at base. Frontal callus 
spade shaped, bulbous, yellow brown, width 
and height subequal. Vertex gray tomentose, 
except glossy brown integument associated 
with each ocellus which extends narrowly to 
upper margin of frontal callus. Mesoscutum 
greenish-gray iridescent, except lateral mar- 
gins yellow tomentose, bearing 3 black lon- 
gitudinal stripes, the median one narrow 
and flanked with gray. Scutellum black ba- 
sally. Pleuron yellow tomentose. Legs 
mostly yellow, fore leg with tarsus and api- 
cal portion of tibia dark brown with black 
hair, mid and hind coxa brown, tarsus with 
black hairs. Wing pattern with broad apical 
spot entering apical half of 2nd submarginal 
cell, apex of hyaline triangle not reaching 
vein R2+3, outer margin of brown crossband 
sinuous, 5th posterior cell infuscated basal- 
ly, hyaline apically, cells br and bm V2 and 
'/s infuscated basally. Halter brown. Tergites 
1-2 yellow brown, tergite 2 with darker 
median integumental marking appearing as 
an inverted "V" set over median yellow- 



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brown triangle, tergites 3-4 predominantly 
brown except for lighter posterior margin 
that expands medianly into light brown pol- 
linose triangle, remaining tergites with an- 
terior % brown, posterior V3 yellow-brown. 
Stemites yellow-brown pollinose with ster- 
nites 3-7 bearing narrow yellow pollinose 
line along the posterior margin, stemites 5- 
7 predominantly brown. 

Male. — Similar to female except for the 
usual sexual differences and following 
characters. Length 6.5-7.5 mm. Pedicel 
only slightly shorter than scape, with finer, 
longer black hairs, length of basal flagel- 
lomere and combined apical flagellomeres 
subequal. Apical spot reaches posterior half 
of 2nd submarginal cell and then fades 
along hind margin, cells br and bm % and 
V2 infuscated basally. First tergite with dark 
spot under scutellum or entirely dark. Fifty 
percent of males examined with abdominal 
pattern of yellow and black with a black 
scutellum. 

Material examined. — 574 9, 30 6, ex- 
amined from the following museums: 
AMNH, CASC, CNCI, CUCC, CUIC, 
FSCA, INHS, TAMU, UGCA, UNHC, 
USNM. 

Distribution. — Massachusetts south to 
Florida, west to Texas; disjunct population 
in Michigan, southern Ontario, and north- 
ern Ohio, Illinois, and Indiana. I have ex- 
amined specimens from the following 
states: AL, CT, DE, FL, GA, IN, KY, LA, 
MA, MD, MI, MS, NC, NH, NJ, NY, SC, 
TN, TX, and VA. L. L. Pechuman has re- 
corded specimens from WV, and Ontario, 
Canada. 

Biology. — Teskey (1969), based his de- 
scription of the larva on 35 specimens. 
They were collected from 3 sites with fresh, 
standing water: wet sand, organic soil, and 
around roots of aquatic vegetation. In 1972, 
Teskey also collected larvae from the edge 
of a slow-moving stream in Virginia (Pe- 
chuman 1973). 

The flight period for Chrysops celatiis is 
late May to September in its northern range, 
and from April to late October in the south. 



Peak flight time over its entire range is in 
June. Although it is a common species, it 
is not abundant enough to be considered a 
pest. 

Discussion. — Pechuman (1949) first de- 
scribed C. celatus as a subspecies of C. 
flovidus, and it was so treated until Teskey 
(1969) found the larvae and pupae of the 
two subspecies to be specifically distinct. 
The adult of C celatus most closely resem- 
bles C. flavidus and C. reicherti, and can be 
separated by its greenish-grey mesoscutum 
and the longer, more slender antennae. The 
sinuous outer margin of the crossband in C. 
celatus is similar to that of C. reicherti, but 
in C. flavidus this margin is either straight 
or concave. The larva resembles C. flavidus 
(Goodwin 1976) and C. dixianus (Teskey 
1969). 

The characters used to define C. celatus 
are consistent, in most cases, but there is 
some variation in the color pattern of the 
abdomen and wings, and in the color of the 
scutellum. The abdominal pattern on ter- 
gites 2-3 may vary from a black inverted 
"V" on the integument to lighter markings 
of dark hairs only. The apical spot of the 
wing may be extended and fade out along 
the hind margin of the wing, approaching a 
pattern much like that of C. reicherti. Spec- 
imens with this wing pattern tend to be 
more robust and have a more darkly infus- 
cated wing. The scutellum, which is usually 
dark basally and reddish apically, is either 
wholly dark or reddish. 

Chrysops dixianus Pechuman 
(Figs. 7, 16, 25) 

Chrysops dixianus Pechuman 1974: 185- 
187. Holotype female: Wedge Plantation, 
McClellanville, South Carolina (CUIC); 
Goodwin 1976: 345-347, description of 
larva and pupa. 

Diagnosis. — Yellow and brown species, 
length 6-8.5 mm. Antenna not swollen, fla- 
gellum V3 longer than scape and pedicel 
combined. Frons width at base subequal to 
width at vertex. Hind coxa and femur dark 



VOLUME 101. NUMBER 1 



185 




atlanticus 




flavidus 




brunneus 




pudicus 




celatus 




reicherti 





dixianus 



sandvi 




tumidicornis 

Figs. 22-30. Female wing patterns. 22. Clinsops atlanticus. 23. C. brunneus. 24, C. celatus. 25, C. dixianus. 
26, C. flavidus. 27, C pudicus. 28. C. reicherti. 29. C. sandyi. 30, C. tumidicornis. 



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brown. Wing picture with 5th posterior cell 
hyaline, infuscated area bordering vein Cu^. 
Female. — Scape and pedicel not notice- 
ably enlarged, yellow with black setae, bas- 
al flagellomere yellow basally, remainder 
brown, apical flagellomeres dark brown to 
black, flagellum approximately V3 longer 
than length of scape and pedicel combined, 
antennal ratio 13:9:32. Frontoclypeus 
glossy yellow, eye margins bearing yellow 
tomentum that extends medianly beneath 
tentorial pits to frontoclypeal suture. Max- 
illary palp light brown with sparse black 
hair basally, more dense apically. Frons 
grayish-yellow pollinose with scattered yel- 
low hairs, width at base subequal to width 
at vertex, height 1.15 times width at base. 
Frontal callus oval, slightly inflated, light 
brown, width 1.4 times height. Vertex yel- 
low gray, glossy surrounding each ocellus, 
ocelli yellow. Mesoscutum yellow-gray to- 
mentose in ground color with 3 dark brown 
longitudinal stripes, median one flanked 
with lighter shade of brown, the two sub- 
lateral stripes merge with median one near 
the scutellum, yellow hair scattered over 
entire mesoscutum. Scutellum dark brown 
basally, light brown apically. Pleuron yel- 
low-gray tomentose. Fore coxa and femur 
yellow, fore tibia with basal V2 yellow, api- 
cal V2 and fore tarsus dark brown, bearing 
black hair. Mid coxa dark brown, femur, 
tibia and metatarsus yellow, remainder of 
tarsus dark brown. Hind coxa and femur 
dark brown, tibia light brown basally, dark- 
ening to apex, metatarsus light brown, re- 
maining portion of tarsus dark brown, tibia 
and tarsus bearing stiff black hair. Wing 
pattern distinct, apical spot broad, filling 
half of 2nd submarginal cell, hyaline tri- 
angle not quite reaching vein R^^^, outer 
margin of crossband straight or convex, 5th 
posterior cell hyaline, vein Cu. infuscated 
along border, cells br and bm V3 and V^ in- 
fuscated basally respectively. Halter with 
stalk brown, knob dark brown. Abdominal 
pattern distinct, tergite 1 yellow, light 
brown under scutellum, tergite 2 with an- 
terior half entirely yellow, posterior half 



yellow with 2 median oblique dark brown 
dashes making a flattened inverted "V" not 
reaching posterior margin of tergite, poste- 
rior margin with median yellow triangle 
and 2 submedian brown spots, tergite 3 
with wide dark brown band that narrows 
medianly, not reaching anterior or posterior 
margins, posterior margin yellow, tergites 
4-5 with anterior V3 dark brown, remaining 
^3 yellow, tergites 6-7 dark brown. Stemites 
1-4 yellow, 4th stemite with dark brown 
median spot, 5th sternite brown with yellow 
hind margin, remaining sternites dark 
brown. 

Male. — Not previously described. 
Length 7.4 mm. Scape and pedicel not no- 
ticeably enlarged, yellow with black hair, 
basal flagellomere yellow at base, remain- 
der brown, apical flagellomeres black, an- 
tennal ratio 15:12:34. Maxillary palp yel- 
low with long yellow hair and a few black 
hairs, length twice width. Mesoscutum yel- 
low-gray tomentose, above wing bases yel- 
low tomentose, with 3 dark brown longi- 
tudinal stripes that merge at the base near 
the scutellum. Scutellum dark brown at 
base, reddish brown apically. Fore coxa and 
femur yellow, tibia with basal Vi yellow, 
apical Vi and tarsus dark brown, bearing 
black hair. Mid coxa dark brown, femur, 
tibia and metatarsus yellow, remainder of 
tarsus dark brown. Hind coxa and femur 
dark brown, tibia light brown basally, dark- 
ening to apex, metatarsus light brown, re- 
maining portion of tarsus dark brown, tibia 
and tarsus bearing stiff black hair. Wing 
pattern with cells br and bm V2 and Va in- 
fuscated basally, hyaline triangle reaching 
vein R2 + 3, infuscated area bordering Cu^ 
vein wider than in female. First abdominal 
tergite yellow, with light brown shading un- 
der scutellum, 2nd tergite with anterior V2 
yellow, posterior V2 with 2 median oblique 
dark brown dashes not attaining posterior 
margin of tergite, posterior margin with 
vague, median yellow triangle, tergite 3 
with wide dark brown median band not 
reaching anterior or posterior margins, bor- 
dered laterally by black setae, hind margin 



VOLUME 101, NUMBER 1 



187 



yellow, tergites 4-5 with anterior % dark 
brown, remaining Vs yellow, tergites 6-7 
dark brown. Sternites 1-3 yellow, the 3rd 
stemite with a dark brown median spot, 
sternites 5-6 brown with yellow hind mar- 
gin, remaining stemite dark brown. Based 
on 1 specimen from Alachua Co. Florida, 
collected by G. B. Fairchild, V-24/25-1975, 
(FSCA). 

Material examined. — 468 9, 1 6, ex- 
amined from the following museums: 
AMNH, CNCI, CUCC, CUIC, FSCA, 
INHS, TAMU, UGCA, UNHC, USNM. 

Distribution. — Virginia to Florida, west 
to Louisiana and Arkansas. I have exam- 
ined specimens from: AL, AR, FL, GA, 
LA, MS, NC, and SC. L. L. Pechuman has 
locality data from VA. 

Biology. — Little is known about the bi- 
ology of Chrysops dixianus. The larva was 
taken from the edge of a freshwater lake in 
mud and decomposing leaves (Goodwin 
1976). Pechuman (1974) stated that C. dix- 
ianus could be a common pest, as suggested 
by 124 specimens collected on July 6, 
1971, in Berkeley County, South Carolina, 
by D. C. Sheppard. 

Discussion. — This species was once con- 
fused with C. pudicus but differs in having 
a longer, thinner flagellum, a brown meso- 
scutum, and the anterior portion of tergite 
2 without markings. The adult of C. dixi- 
anus appears most similar to C. reicherti. 
The 2nd abdominal tergite of both has 
markings only on the posterior Vi, but C 
dixianus is smaller, has a defined apical 
spot, and the 5th posterior cell is hyaline. 

There is little variation in this species. 
The mesoscutum, normally with a narrow 
median brown stripe, may have the stripe 
as wide as the two sublaterals. The scutel- 
lum may be all dark, and the abdominal 
markings on tergites 2-3 can vary from me- 
dian dashes to a more extensive pattern 
reaching the lateral margins. The hind fe- 
mur can be up to Vi light brown. 

Chrysops flavidus Wiedemann 
(Figs. 8, 17, 26) 
Chrysops flavidus Wiedemann 1821: 55. 
Holotype female: Savannah, Georgia 



(MZLU); Krober 1926: 291-292, rede- 
scription; Teskey 1969: 39, description of 
larva and pupa. 

Chrysops pallida Macquart 1838: 162 
(1838: 166). Holotype female: locality 
unknown (MNHN). Synonomized by 
Philip 1965: 325. 

Chrysops pallidus Bellardi 1859: 73. Ho- 
lotype female: Mexico (MRSN). Preoc- 
cupied by Macquart 1838. 

Chrysops guiterasi Brunetti 1923: 401. 
Syntypes female and male: Manzanillo, 
Cuba, 9 (BMNH), 6 (ZMHB). Synon- 
omized by Bequaert, 1940: 279. 

Diagnosis. — Light brown, length 8-10 
mm. Scape and pedicel slightly swollen. 
Mesoscutum yellow with brown stripes. 
Hind margin of crossband straight or con- 
cave, cells br and bm Vi and Vz infuscated 
basally. 

Female. — Scape slightly enlarged, scape 
and pedicel light brown, black setose, basal 
flagellomere light brown, apical flagello- 
meres black, equal to or longer than basal 
flagellomere, antennal ratio 15:10:30. Fron- 
toclypeus glossy yellow, eye maigins bear- 
ing yellow tomentum that extends medianly 
beneath tentorial pits to frontoclypeal su- 
tures. Maxillary palp and labrum yellow. 
Frons predominantly yellow tomentose, 
slightly convergent above, height 1.28 
times width at base. Frontal callus oblong, 
bulbous, hght brown, width 1.83 times 
height. Vertex yellow tomentose, except 
glossy around each ocellus, ocelli black. 
Mesoscutum yellow tomentose, yellow pi- 
lose, bearing three subshiny brown longi- 
tudinal stripes, median one narrower than 
the sublaterals. Scutellum yellow basally, 
orange-yellow apically. Pleuron yellow to- 
mentose. Legs predominantly yellow, mid 
and hind coxa brown, hind tibia with scat- 
tered black hair apically, fore, mid, and 
hind tarsi with black hair. Wing pattern with 
clearly defined hyaline areas, apical spot 
entering posterior half of 2nd submarginal 
cell, apex of hyaline triangle not quite 
reaching vein R2+3, outer margin of cross- 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



band straight or slightly concave, 5th pos- 
terior cell infuscated basally, hyaline api- 
cally, cells br and bm Vi and VS infuscated 
basally. Halter with brown knob and yellow 
stalk. Abdominal tergites golden brown, 
tergites 2-5 with median yellow pollinose 
triangles, each triangle outlined submedi- 
anly with indistinct brown spots, first tergite 
with two submedian brown spots, hind mar- 
gin of tergites 3-6 narrowly yellow tomen- 
tose, tergites bearing mixed black and yel- 
low hairs. Sternites golden yellow tomen- 
tose, sternites 2-6 bearing a narrow yellow 
posterior border, sternites 4-5 with median 
indistinct brown spot. 

Male. — Similar to female except for the 
usual sexual differences and the following 
characters. Length 6.5-7.2 mm. Scape and 
pedicel with longer, finer, black hair. Wing 
faintly tinted below the apical spot and 
along the hind margin, basal Vi of cell br 
infuscated. Abdomen without distinct me- 
dian triangles. 

Material examined. — 1,245 9,15 6 ex- 
amined from the following museums: 
AMNH, BMNH, CASC, CNCI, CUCC, 
CUIC, FMNH, FSCA, INHS, OSUC, 
TAMU, UGCA, UNHC, USNM. 

Distribution. — Missouri to Massachusetts 
south to Texas, Florida, Mexico, the Baha- 
mas, Belize, and Cuba. I have examined 
specimens from AL, AR, DE, FL, GA, IL, 
IN, KS, LA, MA, MD, NC, NJ, NY, OK, 
SC, TN, TX, VA, Belize, the Bahamas, and 
Cuba. L. L. Pechuman recorded specimens 
from CT, lA, KY, MO, MS, OH, PA, RI, 
WV, and Mexico. 

Biology. — The larva described by Teskey 
( 1969) was taken from sand on the edge of 
a pond. Tidwell (1973) collected most of 
the Louisiana specimens from the banks of 
ponds and waterways in bottomland hard- 
wood forests associated with the Mississip- 
pi flood plain. Jones and Bradley (1923) 
reared a larva found at the bottom of a 
brook. Jones and Anthony (1964) collected 
larvae from the margins of brackish water 
and from highly alkaline soil and reared 38 



specimens whose pupal period averaged 8 
days. 

Adults in northern regions, are active 
from June to October and most abundant in 
July. In the south they fly as early as March 
and continue until late September. In some 
areas of Florida there are two distinct pop- 
ulation peaks of Chrysops flavidus, one oc- 
curring in April and May, the other in Au- 
gust and September (Jones and Anthony 
1964). 

This species was described as pestiferous 
to humans (Hine 1906, Jones and Anthony 
1964), generally attacking the head and 
neck, and considered one of the worst 
"stock pests" in the genus Chrysops (Hine 
1906). 

Discussion. — Chrysops flavidus is most 
commonly confused with C atlanticus, C. 
reicherti and C. celatus, all considered at 
one time to be either subspecies or varieties 
of C. flavidus. It is also similar to C. sandyi, 
described below. Chrysops flavidus differs 
from C atlanticus in having thinner anten- 
nae, a yellow mesoscutum, and a wing pat- 
tern with well defined hyaline areas. Chrys- 
ops flavidus differs from C. celatus in its 
shorter, stouter antennae, yellow mesoscu- 
tum, straight or slightly concave outer mar- 
gin of the crossband, and its less distinct 
abdominal pattern. It differs from C reich- 
erti in having the outer margin of the cross- 
band straight or slightly concave, yellow 
hind femur, and an inverted "V" pattern 
that reaches the anterior half of tergite 2. 
Chrysops flavidus differs from C. sandyi in 
its larger size, yellow mesoscutum with 
dark brown stripes, and its more divergent 
frons with the width at the vertex greater 
than the width of the basal callus. 

Variation in C. flavidus is found in the 
tinting of the wing and in the abdominal 
color pattern. Approximately 10% of the 
specimens examined had tinting of the wing 
below the apical spot, approaching the 
smoky wing pattern of C. atlanticus. How- 
ever, C flavidus has a consistently straight 
or slightly concave outer margin of the 
crossband and yellow mesoscutum. Speci- 



VOLUME 101, NUMBER 1 



189 



mens with the apical portion of the wing 
tinted have been collected from the coastal 
areas of South Carolina, Georgia, and Flor- 
ida. A collection of 27 specimens from 
Monroe County, Florida, exhibit a tinted 
wing, have a stouter scape, a grayer meso- 
scutum, and a reddish brown abdomen. 
Other than these differences, these speci- 
mens do not vary in size and other char- 
acters. 

The female syntype of Chrysops guiter- 
asi, described from Cuba and considered a 
synonym of C. flavidus, was examined. I 
have no doubt it is conspecific with C. flav- 
idus, although the abdomen is reddish- 
brown. 

Chrysops piidiciis Osten Sacken 
(Figs. 9, 18, 27) 

Chrysops pudicus Osten Sacken 1875: 381- 
382. Lectotype female: Beverley, Mas- 
sachusetts, (MCZC). Lectotype exam- 
ined, missing head; Goodwin 1976: 350- 
351, description of pupa. 

Diagnosis. — Brown and black, length 
6.5-8.2 mm. Frontal callus reddish-brown 
with black upper margin. Mesoscutum steel 
gray with black stripes. Hyaline triangle 
reaches vein R2 + 3, apical spot narrowly en- 
ters 2nd submarginal cell. Cells br and bm 
Vi and '/f, infuscated basally. 

Female. — Scape and pedicel slightly 
swollen, ground color shiny brown, bearing 
stiff black hairs, basal flagellomere light 
brown gradually darkening at apex, apical 
flagellomeres black, flagellum subequal to 
length of scape and pedicel combined, an- 
tennal ratio 13:9:23. Frontoclypeus glossy 
reddish brown, eye margins bearing light 
brown tomentum that extends medianly be- 
neath tentorial pits to frontoclypeal suture. 
Maxillary palp yellow. Frons light brown 
tomentose, yellow pilose, slightly conver- 
gent above, length 1.3 times width at base. 
Frontal callus oblong, bulbous, reddish 
brown with black upper margin, margin 
concolorous with smooth integument sur- 
rounding ocelli. Vertex dark brown, glossy 



around each ocellus, ocelli black. Scutellum 
black. Fore coxa light brown, trochanter 
slightly darker, femur light brown, apical Va 
darker brown, tibia with black hair apically, 
tarsi black, mid coxa dark brown, femur 
light brown with scattered stiff yellow hair, 
tibia light brown with short black setae api- 
cally, tarsus light brown basally, dark 
brown apically, covered with black setae, 
hind femur dark brown with yellow hair, 
tibia light brown with yellow hair basally, 
remainder with yellow and black hair, tar- 
sus light brown basally, black apically. 
Wing pattern with clearly defined hyaline 
areas, apical spot narrow, entering extreme 
apical portion of 2nd submarginal cell, hy- 
aline triangle reaches vein R2 + 3, outer mar- 
gin of crossband straight or slightly con- 
cave from vein R2+3 to vein M,, cell br and 
bm Vi and /,, infuscated basally. Abdomen 
with first 2 segments yellow, 1st tergite 
with large black spot under scutellum not 
reaching posterior margin, extending later- 
ally to approximately V2 width of tergite, 
2nd tergite with black integumental invert- 
ed "V" not reaching anterior or posterior 
margin, black hair scattered over black in- 
tegumental pattern, orange median triangle 
under black inverted "V" pattern, tergites 
3-4 black anteriorly with median orange 
triangle posteriorly, black pattern not reach- 
ing lateral margin of the tergite, tergites 5- 
6 black anteriorly, yellow posteriorly. Ab- 
domen yellow ventrally, sternites 3-4 with 
median black spot, spot on sternite 3 small- 
er than on sternite 4, remaining sternites 
black. 

Male. — Similar to female except for the 
usual sexual characteristics and the follow- 
ing characters. Scape and pedicel covered 
with long, fine, black hair. Mesoscutum al- 
most entirely black, the 3 stripes indistinct, 
with scattered yellow hair. Apical spot 
slightly more extensive. Cell br % infuscat- 
ed basally. 

Material examined. — 91 9, 1 d exam- 
ined from the following collections: CUIC, 
FSCA, UNHC. 

Distribution. — Southern Illinois to Mas- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



sachusetts, south to eastern Texas and Flor- 
ida with some disjunct populations in north- 
em Indiana, Michigan, New York, southern 
Ontario, and Nova Scotia. I have examined 
specimens from AL, CT, FL, GA, LA, LA, 
NC, NH, NY, RI, SC, and Nova Scotia. L. 
L. Pechuman has records from NJ, DE, IN, 
IL, MD, MI, MS, OK, TN, TX, and WI. 

Biology. — The larva of this species, al- 
though not formally described, has report- 
edly been taken from wet soil along a road- 
side ditch (Jones and Anthony 1964, Pe- 
chuman 1973). The pupa was described by 
Goodwin (1976) from Baldwin County, Al- 
abama, but the habitat is unknown. Jones 
(1953) indicated that in parts of Florida 
there are two generations of C. piidicus an- 
nually, one that emerges in May-June and 
another in August-September. Large num- 
bers can occur in wooded areas of Florida 
during April and May (Jones and Anthony 
1964). 

Discussion. — Pechuman et al. (1983) 
treated C. pudicus as a member of the 
Chrysops callidus group, which includes C. 
dimmocki Hine, a species that can resemble 
C. pudicus. Members of the C. callidus 
group all have a black frontal callus, in- 
verted "V" pattern on 2nd abdominal ter- 
gite that reaches the anterior margin and, in 
some species, a hyaline triangle that ex- 
tends beyond vein R2+3. Chrysops pudicus 
differs from members of the C callidus 
group in having an abdominal pattern on 
tergite 2 not reaching the anterior margin, 
a hyaline triangle not crossing vein R2+3. 
and a yellow frontal callus with dark top 
margin. The frontal callus is rarely entirely 
dark (Brennan 1935, Teskey 1990), and the 
hyaline triangle rarely reaches beyond vein 
R2+3. Therefore, I have chosen to place C. 
pudicus in the Chrysops flavidus group. 

Chrysops pudicus most closely resembles 
C. tumidicornis, described below, in having 
similar size, dark coloration, and 1st ab- 
dominal tergite with a dark spot under the 
scutellum. They differ in that C. pudicus 
has a narrower scape and pedicel, a less ex- 



tensive, well defined apical spot, and a hy- 
aline triangle that reaches R2+3. 

Variation within C. pudicus occurs in the 
extent of the hyaline triangle (Pechuman 
1973) and in the color of the frontal callus. 
Of the 91 females examined, 10 specimens 
from Florida had a hyaline triangle that did 
not reach vein R2+3. Although variation in 
color of the frontal callus is mentioned in 
various publications (Brennan 1935, Teskey 
1990), all the specimens I examined had a 
yellow frontal callus with a dark upper mar- 
gin. 

Chrysops reicherti Fairchild 
(Figs. 10, 19, 28) 

Chrysops reicherti Fairchild 1937: 60-61. 
Holotype female: Monticello, Jefferson 
County, Florida (MCZC). Holotype ex- 
amined. Goodwin 1972: 105-107, de- 
scription of larva and pupa. 

Chrysops flavida reicherti: Philip, 1947: 
273, treated as a subspecies. Pechuman 
1957: 30-31, description of male; Philip 
1965: 325, treated as a variety of C. flav- 
idus. 

Diagnosis. — Yellow and brown, length 
8-9.5 mm. Antenna not swollen. Meso- 
scutum yellow in ground color with 3 
brown stripes. Tergites 1-2 light yellow, 
darker pattern on posterior half of tergite 2. 
Cells br and bm are V-i and Va infuscated 
basally. 

Female. — Scape and pedicel not swollen, 
light brown with black setae, flagellum 
black, antennal ratio 15:10:35. Frontocly- 
peus glossy yellow, eye margins bearing 
grayish yellow tomentum extending medi- 
anly beneath tentorial pits to frontoclypeal 
suture. Maxillary palp brown. Frons yellow 
tomentose, slightly convergent above, 
height 1.3 times width at base. Frontal cal- 
lus oblong, bulbous, light brown, width 
1.33 times height. Vertex light brown 
around each ocellus, ocelli yellow. Meso- 
scutum yellow tomentose with three sub- 
shiny brown longitudinal stripes, median 
one narrower than sublaterals, flanked with 



VOLUME 101, NUMBER 1 



191 



lighter shade of brown. Scutellum yellow 
brown. Pleuron yellow tomentose. Legs 
predominantly light brown, fore and mid 
tarsi brown, mid and hind tibiae with dark 
hair apically, hind femur reddish brown, 
apical tarsomeres brown to black. Wing 
with extensive apical spot reaching poste- 
rior Vi of 2nd submarginal cell fading into 
3rd posterior cell, hyaline triangle not 
reaching vein R2 + 3, outer margin of cross- 
band bowed outward at center, 5th posterior 
cell infuscated basally and along vein mar- 
gins, hyaline apically, cells br and bm V3 
and Va infuscated basally. Halter with brown 
stalk and knob. First abdominal tergite light 
brown, second tergite with basal half yel- 
low, green tinted medianly, apical half with 
oblique brown spots reaching hind margin 
and enclosing pale triangle. Tergite 3 with 
brown median band set over median orange 
triangle, hind margin of tergite gray. Ter- 
gites 4-6 with basal half brown, apical half 
yellow. Sternites predominantly golden 
brown, 3-6 with median brown spot pro- 
gressively enlarged on posterior segments. 

Male. — Similar to female except for the 
usual sexual differences and the following 
characters: Length 6.5-7.0 mm. Scape 
swollen, pedicel V^ longer than in female, 
scape and pedicel with longer, finer, black 
hair. Mesoscutum with long yellow hair. 
Wing with apical spot more extensive, cells 
br and bm V2 and Vs infuscated basally. 

Material examined. — 720 9 , 20 d ex- 
amined from the following collections: 
CASC, CNCL CUIC, FSCA, INHS, 
TAMU, UGCA, UNHC, USNM. 

Distribution. — Southern Illinois to Dela- 
ware, south to eastern Texas and Florida. I 
have examined specimens from AL, AR, 
FL, GA, IL, KY, LA, MA, MD, MS, NC, 
SC, TN, TX, and VA. L. L. Pechuman has 
records from IN and MO. 

Biology. — The larvae of Chrysops reich- 
erti have been collected in very wet mud 
and organic debris from small ponds and 
lakes (Goodwin 1972, Tidwell 1973). 
Adults have been collected between April 
and September in the southern states. The 



one Massachusetts specimen was collected 
on May 8. 

Discussion. — Chrysops reicherti was 
originally described as a species by Fair- 
child (1937). Philip (1947) listed it as a sub- 
species of C. flavida. Goodwin (1972), after 
describing the larva and pupa of C. reich- 
erti and comparing them to those of C. flav- 
idus, considered them separate species and 
elevated C. reicherti to a full species. 

Chrysops reicherti is most commonly 
confused with C flavidus, but is separated 
by its pale yellow first two abdominal seg- 
ments, sinuous outer margin of the cross- 
band, and the slightly darker hind femora. 
In the key above, C reicherti shares with 
C dixiamis the color pattern on the poste- 
rior portion of the second tergite. They dif- 
fer in that C. dixianus has a straighter outer 
margin of the crossband, an apical spot that 
reaches slightly beyond the apex of the 
wing, and 5th posterior cell extensively hy- 
aline. 

Variation in C reicherti includes tinting 
in the usual hyaline areas of the wing, 2nd 
abdominal tergite without the greenish cast 
described by Fairchild and, hind femur light 
or dark brown. 

Chrysops sandyi Baier, new species 

(Figs. 12, 20, 29) 

Diagnosis. — Yellow and brown, length 
6-8 mm. Antenna slightly enlarged. Me- 
soscutum with brown iridescent stripes. 
Wing slightly tinted in usual hyaline areas. 
Cells br and bm are Vi and V3 infuscated 
basally. 

Holotype female. — Length 6.2 mm. 
Scape slightly enlarged, antenna light 
brown except apical 3/4 of 1st flagellomere 
slightly darker, apical 4 flagellomeres black, 
antennal ratio 13:9:24, length of flagellum 
subequal to combined length of scape and 
pedicel. Frontoclypeus glossy yellow, eye 
margins bearing yellow tomentum that ex- 
tends medianly beneath tentorial pits to 
frontoclypeal suture. Maxillary palp light 
brown with sparse black hairs, length 4 
times width. Frons yellow tomentose with 



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scattered yellow hairs, convergent above, 
height slightly more than width at base, 
width at vertex less than width of frontal 
callus. Frontal callus oval, yellow brown, 
width 1.7 times height. Vertex yellow to- 
mentose, glossy around each ocellus, ocelli 
light brown. Mesoscutum yellow-green iri- 
descent tomentose, except lateral margins 
iridescent yellow gray, bearing 3 brown to 
dark brown stripes, the median one darker 
and narrower than the two sublaterals. Scu- 
tellum dark brown basally, brown apically. 
Pleuron yellow gray. Legs mostly light 
brown, fore tarsus black, apical portion of 
tibia dark brown with black hair, mid coxa 
dark brown, tarsus with black hair, hind 
coxa dark brown, femur and basal tibia red- 
dish brown, apical tibia brown, apical tibia 
and tarsus with black hair. Wing with broad 
apical spot reaching posterior half of 2nd 
submarginal cell, apex of hyaline triangle 
including posterior half of 1st submarginal 
cell, not reaching vein R2+3, outer margin 
of crossband straight, 5th posterior cell 
slightly tinted, infuscated along vein Cu2, 
cells br and bm Vz and V3 infuscated basally. 
Halter with light brown stalk and brown 
knob. Tergites 1-2 yellow brown, tergite 1 
with a few dark hairs under scutellum, ter- 
gite 2 with a median brown integumental 
marking appearing as a flattened inverted 
"V", not reaching anterior margin, fading 
into pale posterior margin, median orange- 
brown triangle behind inverted "V pat- 
tern, tergite 3 mottled dark brown anteri- 
orly, orange brown posteriorly, extending 
medianly to form orange triangle, tergites 
4-6 dark brown anteriorly, yellow orange 
posteriorly, tergites 3-6 with gray hind 
margin. Sternites 1-3 yellow brown, ster- 
nite 3 bearing a median brown spot, ster- 
nites 4-7 dark brown, posterior margin pale 
yellow tomentose. 

Male. — Unknown 

Material examined. — Holotype $, AL- 
ABAMA: Mobile County, VII- 1952, T. R. 
Adkins (CUIC). Paratypes: 105 ?. ALA- 
BAMA: Mobile County, VII- 1952 (62 $ ), 
VII-24-1952 (4 9), T. R. Adkins (CUIC); 1 



9 Corden, VII-8-1962, M. Tidwell (FSCA). 
LOUISIANA: St. Tammany Parish, 4 $ 
Slidell, 7 mi SE,VI-23-1980, E. A. Lisow- 
ski (CUIC). MISSISSIPPI: Hancock Coun- 
ty, Ansley, VI- 10- 1966 (5 9), VI-20-1966 
(6 9), Diamond & Bradford (FSCA); 2 9 
V-5-1976, M. Tidwell (FSCA); 1 9 Pear- 
lington, VI-9-1966, ? collector (FSCA); 1 
9 MTO fee area, IV-27-1965, R. Hepburn 
(FSCA). Jackson County, Fountainbleau, 
VI-26-1966 (2 9), VII-27-1968 (1 9), B. 
Byrd (FSCA); 4 9 Gautler, 4.5 mi WSW, 
VI-24-1980, E. A. Lisowski (CUIC); 12 9 
Ocean Springs, V-10-1970, G. Ross 
(FSCA). 

Etymology. — Named in honor of G. B. 
"Sandy" Fairchild for his extensive contri- 
bution to the organization of Chrysops taxa 
and his desire to see the southeastern spe- 
cies clarified. His examination of specimens 
and critical comments on this work has 
been greatly appreciated. 

Distribution. — Gulf coast of Alabama, 
Mississippi, and Louisiana. 

Biology. — Chrysops saudyi has been col- 
lected between April 27 and August 8. 
Based on its distribution along the Gulf 
coast, it is probably limited to a litoral en- 
vironment. 

Discussion. — This species has been con- 
fused with C. atlauticus because of the tint- 
ed wing and greenish iridescent color of the 
mesoscutum. Chrysops sandyi has a nar- 
rower antennae, the pedicel % the length of 
the scape, longer palpi, length 4 times width 
of apical palpomere, mesoscutum with 
brown longitudinal stripes, and lightly tint- 
ed wing pattern. Chrysops atlanticus inhab- 
its the Atlantic coast from Maine to Florida 
while C. sandyi is found along the Gulf 
coast of Alabama, Mississippi, and Louisi- 
ana. 

Variation in C. sandyi occurs in the 
ground color of the mesoscutum, and in the 
abdominal pattern. Ground color of the me- 
soscutum, although usually greenish irides- 
cent, may have yellow tomentum. Abdom- 
inal pattern may be indistinct, with a pattern 
of black hair and no integumental markings. 



VOLUME 101. NUMBER 1 



193 



Specimens with these features appear sim- 
ilar to C. flavidus, but C sandyi is smaller, 
wing tinted in the usual hyaline areas, and 
flagellum subequal to the combined length 
of the scape and pedicel. 

Chrysops tumidicornis Baier, 
new species 

(Figs. 11, 21, 29) 

Diagnosis. — Black and brown, length 
5.8-8 mm. Antenna swollen, pedicel barrel 
shaped. Frontal callus width approximately 
2 times height. Mesoscutum grey green 
with black stripes. Wing with distinct hya- 
line triangle, cells br and bm % and Va in- 
fuscated basally. 

Holotype female. — Length 7.4 mm. 
Scape and pedicel swollen, pedicel barrel 
shaped, brown, covered with black setae, 
basal flagellomere brown, apical 4 flagel- 
lomeres black, flagellum subequal to length 
of scape and pedicel combined, basal fla- 
gellomere equal to combined length of the 
apical 4 flagellomeres, antennal ratio 14:1 1: 
26. Frontoclypeus glossy yellow, eye mar- 
gins bearing yellow tomentum that extends 
medianly beneath tentorial pits to frontocly- 
peal suture. Maxillary palp light brown 
with scattered stiff black hairs, length 4 
times width of apical palpomere. Frons yel- 
low tomentose, slightly convergent above, 
height 1.2 times width at base. Frontal cal- 
lus oval, slightly bulbous, yellow, width 1 .9 
times height. Vertex yellow brown except 
glossy black around each ocellus, ocelli 
black. Mesoscutum grayish-green ground 
color with 3 longitudinal black stripes not 
merging basally, median one narrower than 
two sublaterals, yellow tomentose above 
wing bases. Scutellum black basally, red- 
dish brown apically. Fore coxa, femur and 
basal V-i of tibia yellow, remaining tibia 
brown, black setose, tarsus entirely dark 
brown, black setae, mid coxa brown, femur, 
tibia and basal tarsomeres yellow, tarsi and 
apical Va of tibia with black setae, hind 
coxa, femur and tibia brown, scattered yel- 
low hair and black setae, black setae be- 
coming more dense toward apical portion 



of tibia and tarsus. Wing pattern distinct, 
apical spot includes apical % of vein R4, 
reaching posterior half of 2nd submarginal 
cell, and fading along hind margin of wing 
to crossband, hyaline triangle slightly tint- 
ed, not reaching vein R2+3, outer margin of 
crossband straight, 5th posterior cell infus- 
cated along margins, center hyaline, cells br 
and bm % and Va infuscated basally respec- 
tively. Abdominal tergites orange yellow, 
1st tergite dark brown under scutellum, ter- 
gite 2 with median black inverted "V" pat- 
tern on integument not reaching anterior, 
posterior or lateral margins, scattered black 
hair over dark pattern, median orange tri- 
angle behind inverted "V" pattern, hind 
margin gray, tergite 3 with gray posterior 
border widened medianly to form an indis- 
tinct triangle, anterior half with black pat- 
tern of 2 half circles that merge above tri- 
angle and occupy V^. width of tergite, ter- 
gites 4-6 black anteriorly, posterior border 
gray. Abdomen orange yellow ventrally, 
sternites 3-5 with median dark spot becom- 
ing wider on each successive sternite, ster- 
nites 6-7 dark brown. 

Male. — Unknown. 

Material examined. — Holotype 9, 
FLORIDA: Levy County, Cedar Key Shell 
Mound, VII-7-1976, R. H. Roberts (FSCA). 
Paratypes: 206 ?. FLORIDA: Bay County, 
2 9 V- 14- 1957, F W Mead (CASC); 1 9 
V-25-1982, J. Hogsette (FSCA); Saint An- 
drews St. Park, 1 9 V-14-1957, F W Mead 
(CASC). Citrus County, 3 9 Ozello, VII- 
29-1959, H. V. Weems (CASC). Collier 
County, 1 9 Collier-Seminole State Park, 
XI-26-1976, John Edward Rawlins (CUIC). 
Dade County, 1 9 Everglades National 
Park, 1-29-1959, H. A. Denmark (CASC); 
1 9 Royal Palm Hammock, VI-22-1951, 
Price, Beamer & Wood (CASC); 1 9 West 
Lake, XII-4-1970, P H. & M. Arnaud 
(CASC). Dixie County, 2 9 Jena, VII-27- 
1991, Jena, 10 mi S on RT 361, VII-22- 
1985 (2 9), VII-13-1991 (8 9), VII-21- 
1991 (5 9), VII-27-1991 (4 9), VIII-31- 
1991 (13 9), VII-1 1-1992 (1 9) L. R. Da- 
vis, Jr. (FSCA); 3 9 Suwannee, V- 19- 1964, 



194 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



C. F. Zeiger (CASC); 2 9 Suwannee, V-19- 
1964 (FSCA). Franklin County, Carrabelle, 
IV-25-1980 (2 ?), IV-27-1980 (3 $ ), IV- 
28-1980(1 9 ), I V-29- 1980(4 9), V-4- 1 980 
(5 9), IV- 14- 1977 (1 9), L. L. Pechuman 
(CUIC); I V-28- 1980(1 9), V-4- 1980 (1 9), 
L. L. Pechuman (FSCA); Eastpoint, 1 9 V- 
19-1935 (FSCA); Lake Morality, IV-27- 
1980 (1 9), V-10-1980 (1 9), 1 9 Timber 
Island, V-4- 1980, L. L. Pechuman (CUIC); 

1 9 Timber Island, V-4- 1980, L. L. Pechu- 
man (CASC); 2 9 St. George Island, IV-5- 
1976, L. L. Pechuman (FSCA). Gilchrist 
County, 1 9 VI-2-1950, (CASC). Gulf 
County, 4 9 V-1 1-1973, J. T. Goodwin 
(FSCA), V-4- 1973 (19), VIII- 14- 1971 (1 
9), H. V Weems, Jr (FSCA). 2 9 St. Jo- 
seph State Park, V- 1/3- 1970, W. W. Wirth 
(USNM); 1 9 V-5-1987, L. Strange and J. 
Wiley (FSCA). Hernando County, 1 9 Bay- 
port, IV-22-1978, L. A. Strange (FSCA). 
Hillsborough County, 1 9 Tampa, IV-29- 
1950, G. B. Worth (CASC). Jefferson 
County, 1 9 Monticello, IX-18-1935, G. B. 
Fairchild (FSCA). Lee County, 2 9 Sanibel 
Island, IV-8-1933, W. J. Clench (USNM); 2 
9 Sanibel Island IV-8-1933, W J. Clench 
(FSCA). Levy County, 2 9 VII-8-1980, E. 
Davis (FSCA); 1 9 V-6-1955, H. V. Weems 
(FSCA); 3 9 Cedar Key Vl-20-1974, 
(FSCA); 5 9 Cedar Key area, VI-4-1991, 
J. Huether (CUIC); 3 9 Cedar Key, VII- 12- 
1939, R. H. Beamer (CASC); 3 9 VI- 12- 
1939, Oman (USNM); 1 9 VII-29-1977, L. 
R. Davis, Jr (FSCA); 1 9 VI-28-1973, Carl 
Shleck (FSCA); 1 9 VI-15-1979, L. A. 
Wood (FSCA); Cedar Key, Shell Mound, 
VI-9-1971 (5 9), VI- 16- 1973 (1 9 ), VI-29- 
1976 (6 9), VII-3-1976 (7 9), G. B. Fair- 
child (FSCA); 1 9 3 mile E. Shell Mound, 

2 9 VI-9-1971, (FSCA); 8 9 VII-7-1976, 
R. Roberts (FSCA); 3 9 VI-21-1973, R. 
Wilkerson (FSCA); Yankeetown, V-8/13- 
1980(1 9), VII-8-1980 (1 9), V-16-1980 
(3 9), VI-24-1980 (5 9), E. Davis (FSCA); 
V-2 1/23- 1979 (19), V- 14/1 6- 1979 (2 9), 
V-1 1/14-1979 (19), V-23/24-1979 (3 9), 
V-25/30-1979 (2 9), V-30/3 1-1979 (19), 
Roberts (FSCA); 3 9 V-30/3 1-1985, D. L. 



Kline (FSCA); VIII- 10- 1966 (1 9), VII-28- 
1965 (4 9), C. F Zeiger (FSCA). Manatee 
County, 1 9 VI- 12- 1925, T H. Hubbell 
(CASC). Monroe County, 1 9 Everglades 
National Park, IV-8-1970, W. W. Wirth 
(USNM). Okaloosa County, 4 9 V-12- 
1972, J. T. Goodwin (FSCA). Pasco Coun- 
ty, 5 9 Hudson, VII- 13- 1939, R. H. Beamer 
(CASC); 11 9 VII-13-1939, Oman 
(USNM); 1 9 VII-13-1939, Oman (FSCA); 
2 9 VII-13-1939, D. E. Hardy (CASC): Pi- 
nellas County, 1 9 IV- 19- 1930, B. R Moora 
(USNM); 1 9 V-2-1955, R. R Essar 
(FSCA); 1 9 V-1 0-1 955, E. W. Holder, Jr. 
(FSCA); 1 9 Dunedin, V- 14- 1959, O. L. 
Cartwright (USNM); 2 9 Pass-a-grille, IV- 
7-1930, W. G. Fargo (CASC). Santa Rosa 
County, 1 9 Santa Rosa Island, V-24-1971, 
H. V. Weems Jr. (FSCA); Taylor County, 1 
9 Steinhatchee, VII- 12- 1966, R. R Esser 
(CASC); 1 9 Steinhatchee, VII-20- 1991, L. 
R. Davis, Jr. (FSCA); 1 9 12 mi NW of 
Steinhatchee, V-24-1983, R. M. Reeves 
(UNHC); 12 9 Cedar Island, 16 miles NW 
of Steinhatchee, V-1 6- 1969, H. V. Weems, 
Jr. (FSCA); 3 9 Cedar Island. 16 miles NW 
of Steinhatchee, V-1 6- 1969, H. V. Weems, 
Jr. (CASC). Wakulla County, 1 9 V-2- 
1980, L. L. Pechuman (CUIC); 3 9 Och- 
lockonee River State Park, V-20-1968, H. 
V. Weems, Jr. (FSCA); 2 9 Ochlockonee 
River State Park, V-20-1968, H. V. Weems, 
Jr. (CASC); 1 9 Ochlockonee River State 
Park, VI-22-1973, Fairchild & Wilkerson 
(FSCA). 

Etymology. — This species is named for 
its characteristic "robust" pedicel. 

Distribution. — West coast of Florida. 

Biology. — Based of the collection data, 
most adults have been taken between 
April-August with a few collected in Sep- 
tember, and November-January. All speci- 
mens examined were taken from coastal 
counties near brackish habitats. Although 
the larva and pupa are unknown, I suspect 
this species is limited to salt marshes. 

Discussion. — This species has been con- 
fused with C. atUmticus because they both 
have swollen antennae. Also, I believe that 



VOLUME 101, NUMBER 1 



195 



Pechuman (1949), in his description of C. 
atlanticus, had before him specimens of this 
species. He wrote that many specimens ex- 
amined showed considerable melanism, 
varying from slight enlargement of abdom- 
inal spots to extreme enlargement where the 
tergites were mostly dark with a narrow 
yellow-brown margin, a pattern I find to be 
diagnostic C. tumidicornis. 

In the commonly used keys to species of 
North American Chrysops (Jones and An- 
thony 1964, Teskey 1969, Tidwell 1973, 
Pechuman 1973, 1981) C tumidicornis wiW 
key to C. atlanticus. However, C. tumidi- 
cornis differs by having a stouter pedicel, 
frontal callus approximately twice as wide 
as tall, wing pattern with a distinct hyaline 
triangle, and an abdominal pattern brown 
with median orange triangles. Chrysops 
tumidicornis has an extensive apical spot 
reaching posterior half of 2nd submarginal 
cell, and a hyaline triangle that does not 
reach vein Ri^- Chrysops tumidicornis ap- 
pears to be more similar to C pudicus in 
size, dark mesoscutum and abdominal pat- 
tern. 

Variation within C. tumidicornis ranges 
from specimes with small, slender body, 
and a more uniformly dark abdomen, to a 
larger, more robust body, with an abdomi- 
nal pattern more uniformly yellow-orange. 
Three specimens from Gulf County, Flori- 
da, have an entirely dark brown abdomen 
with a lighter median triangle on the 2nd 
tergite. Chrysops tumidicornis is easily rec- 
ognized by its bulbous, barrel shaped ped- 
icel. 

Acknowledgments 

I acknowledge the assistance of the late 
G. B. "Sandy" Fairchild, Florida State Col- 
lection of Arthropods, for assistance with 
organization of data and critical comments 
on treatment of species, the late L. L. Pe- 
chuman, whose distribution records were 
used to create accurate distribution maps for 
this paper, John F. Burger, University of 
New Hampshire, for providing critical com- 
ments, financial support and encouragement 



throughout this project, Donald S. Chan- 
dler, University of New Hampshire, for his 
assistance with development of the data- 
base, illustrations and editorial comments, 
the late R. Marcel Reeves, University of 
New Hampshire, for editorial comments, 
John Weaver, University of New Hamp- 
shire, for assistance with illustrations, and 
finally the individuals responsible for lend- 
ing specimens held by their respective in- 
stitutions. 

This is scientific contribution number 
1967 from the New Hampshire Agriculture 
Experiment Station. 

Literature Cited 

Anderson. J. F. 1971. Autogeny and mating and their 
relationship to biting in the saltmarsh deer fly, 
Chrysops atlanticus (Diptera: Tabanidae). Annals 
of the Entomological Society of America 64: 
1421-1424. 

Arnett, R. H., Samuelson, G. A., and Nishda, G. M. 
1993. The insect and spider collections of the 
world. Flora and Fauna Handbook No. 1 1 . 306 pp. 

Bellardi, L. 1859. Saggio di Ditterologia Messicana. 
Memorie della Reale Academia dclle Science di 
Torino 19: 246-276. 

Bequaert, J. 1940. The Tabanidae of the Antilles 
(Dipt.) Revista de Entomologia 1 1: 279. 

Brennan, J. M. 1935. The Pangoniinae of Nearctic 
America (Diptera: Tabanidae). The University of 
Kansas Science Bulletin 22(13): 1-400. 

Brunetti, E. 1923. Two new species of Tabanidae from 
Cuba. Bulletin of Entomological Research 13: 
401-402. 

Burger, J. E 1995. Catalog of Tabanidae (Diptera) of 
North America north of Mexico. Contributions to 
Entomology, International 1(1): I-IOO. 

Fairchild, G. B. 1937. A preliminary list of the Taban- 
idae (Diptera) of Florida. Florida Entomologist 
19: 58-63. 

Goodwin, J. T. 1972. Immature stages of some eastern 
nearctic Tabanidae (Diptera). I. Introduction and 
the genus Chrysops Meigen. Journal of the Geor- 
gia Entomological Society 7(2): 98-109. 

. 1976. Immature stages of some eastern nearc- 
tic Tabanidae (Diptera). VI. Additional species of 
Chrysops Meigen. Florida Entomologist 59(4): 
343-35 1 . 

Hansens, E. J. 1980. Review: Tabanidae of the east 
coast as an economic problem. Bulletin of the 
New York Entomological Society 137(4): 312- 
318. 

Hine, J. S. 1903. Tabanidae of Ohio with a catalogue 
and bibliography of the species from America 



196 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



north of Mexico. Ohio State Academy of Science, 
Special Papers 5: 1-63. 
. 1906. A preliminary report on the horseflies 



of Louisiana with a discussion of remedies and 
natural enemies. State Crop Pest Commission of 
Louisiana. Circular No. 6. 43 pp. 

International Commission on Zoological Nomencla- 
ture. 1993. Opinion 1711. Chnsops atlanticiis Pe- 
chuman. 1949 (Insecta, Diptera): specific name 
conserved. Bulletin of Zoological Nomenclature 
50: 83. 

Jamnback, H. and Wall. W. 1959. The common salt- 
marsh Tabanidae of Long Island, New York. New 
York State Museum and Science Service Bulletin 
Number 375. 1-72. 

Jones, C. M. 1953. Biology of Tabanidae in Florida. 
Journal of Economic Entomology 46(6): 1108- 
II09. 

Jones, C. M., and Anthony, D. W. 1964. The Tabanidae 
(Diptera) of Florida. United States Department of 
Agriculture Technical Bulletin 1295, 85 pp. 

Jones, T. H. and Bradley W. G. 1923. Observations on 
Tabanidae (horseflies) in Louisianna. Journal of 
Economic Entomology 16: 307-312. 

Krober, O. 1926. Die Chrysops Arten Nordamerikas 
einshl. Mexicos. Stettiner Entomologische Zei- 
tung 87: 1-353. 

Macquart, P. J. 1838. Dipteres Exotiques. Nouveaux 
peu connus. Part I. Tabaniens. Librairie Encyclo- 
pedique de Roret, Paris. Pages 90-164. 

Magnarelli. L. A. and Anderson, J. E 1976. Follicular 
development in salt marsh Tabanidae (Diptera) 
and incidence of nectar feeding with relation to 
gonotrophic activity. Annals of the Entomological 
Society of America 70: 529-533. 

Orihel, T C. and Lourie, R. C. 1975. Loa loa: devel- 
opment to the infective stage in an American deer- 
fly, Chrysops atlanticiis. American Journal of 
Tropical Medicine and Hygiene 24(4): 610-615. 

Osten Sacken, C. R. 1875. Prodrome of a monograph 
of the Tabanidae of the United States. Part I. The 
genera Pangonia. Chrysops, Silviiis, Haematopo- 
ta. Diahasis. Memoirs of the Boston Society of 
Natural History 2: 381-382. 

Pechuman, L. L. 1949. Some notes on Tabanidae (Dip- 



tera) and the description of two new Chrysops. 

The Canadian Entomologist 131(3): 77-84. 
. 1957. Descriptions of Tabanidae previously 

known from one sex only (Diptera). Bulletin of 

the Brooklyn Entomological Society 102(2): 29- 

34. 
. 1973. Horse flies and deer flies of Virginia 

(Diptera: Tabanidae). The Insects of Virginia No. 

6. 92 pp. 
. 1974. Two new Tabanidae from southeastern 

United States (Diptera). Journal of the New York 

Entomological Society 82(3): 183-188. 
. 1981. The horse flies and deer flies of New 



York (Diptera: Tabanidae). Search Agriculture. 
Cornell LIniversity Agriculture Experiment Sta- 
tion. (Ithaca, N.Y) 2: 1-65. 

Pechuman, L. L., Webb, D. W., and Teskey, H. J. 1983. 
The Diptera, or True Flies, of Illinois, I. Tabani- 
dae. Illinois Natural History Survey Bulletin 
33(1): 1-120. 

Philip, C. B. 1947. A catalog of the blood-sucking fly 
family Tabanidae (horseflies and deerflies) of the 
Nearctic Region North of Mexico. The American 
Midland Naturalist 37(2): 257-324. 

. 1955. New North American Tabanidae. IX. 

Notes on and keys to the genus Chiysops Meigen. 
Revista Brasileira del Entomologia 3: 47-128. 

Philip, C. B. 1965. Family Tabanidae, pp. 319-342. //; 
Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. 
Foote, and J. R. Coulson, eds., A Catalog of the 
Diptera of America North of Mexico. U.S. De- 
partment of Agiculture. Agricultural Research 
Service Handbook 276, 1696 pp. 

Teskey, H. J. 1969. Larvae and pupae of some eastern 
North American Tabanidae (Diptera). The Ento- 
mological Society of Canada 63: 1-147. 

. 1990. The horse flies and deer flies of Canada 

and Alaska (Diptera: Tabanidae). The Insects and 
Arachnids of Canada. Biosystematics Research 
Institute, Ottawa 16: 1-381. 

Tidwell. M. A. 1973. The Tabanidae (Diptera) of Lou- 
isiana. Tulane Studies in Zoology and Botany 
18(1-2): 1-95. 

Walker, F 1848. List of the specimens of dipterous 
insects in the collection of the British museum 1 : 
229 pp. London. 

Wiedemann, C. R. W. 1821. Diptera exotica (ed. 2), 
1-244, 1 fig., 2 plates. Kiliae (= Kiel). 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999. pp. 197-207 

TWO NEW SPECIES OF PHAENOCARPA FOERSTER (HYMENOPTERA: 
BRACONIDAE: ALYSIINAE) FROM SOUTH AMERICA 

M. Trostle, N. S. Carrejo, I. Mercado, and R. A. Wharton 

(MT, IM, RAW) Department of Entomology, Texas A&M University, College Station, 
TX 77843, U.S.A. (e-mail: RAW: rawbaw2@acs.tamu.edu); (NSC) Departamento de Bio- 
logia, Seccion Entomologia, Universidad del Valle, A. A. 25360, Cali, Colombia 



Abstract. — Two new species of Phaenocorpa Foerster are described from South Amer- 
ica: P. hyalina Trostle and P. pericarpa Wharton and Carrejo. Phaenocarpa pericarpa 
was reared from Anastrepha distincta Greene in pods of Inga, representing the first host 
record for Phaenocarpa in South America. A key separating the South American species 
of Phaenocarpa is included. 

Key Words: Anastrepha; parasitoid; Diptera; Neotropical 



Papp (1969) revised the species of 
Phaenocarpa Foerster for the Neotropical 
Region, treating 10 species. Papp (1966, 
1969) included Asobara Foerster as a syn- 
onym of Phaenocarpa in his earlier works, 
but Fischer (1971) subsequently removed 
Asobara from synonymy and it has been 
recognized as a separate genus ever since. 
Consequently, five of the 10 species treated 
by Papp (1969) must now be placed in A^o- 
bara. These are anastrephae (Muesebeck), 
gahani (Papp), mexicana ( Ashmead), p/^nr- 
alis (Ashmead), and rubra (Papp). Their 
identity has been confirmed by one of us 
(RAW), and the placement of two of these 
species has been discussed previously 
(Wharton 1994). 

A sixth species, delicata Papp, has sev- 
eral unusual features, and its placement is 
uncertain (Wharton 1980, Fischer 1994). 
Papp (1969) divided the four remaining 
species into those with well-developed no- 
tauli and those without. Wharton (1994) de- 
scribed three additional species from the 
Neotropics. He further defined two distinct 
species groups to accomodate these, and 
discussed the differences which set these 



two groups apart from other described spe- 
cies of Phaenocarpa. 

Two more species are described here. 
The first belongs to the cratomorpha Whar- 
ton species group as defined by Wharton 
(1994). The second belongs to Papp's 
(1969) group with reduced notauli. 

The genus Phaenocarpa is fairly large, 
but hosts have been recorded for only about 
15% of the 150 described species (Fischer 
1974, 1975, 1990, 1993; Shenefelt 1974; 
Wharton 1984; Vet and van Alphen 1985; 
Tobias 1986). Holarctic species have been 
reared most frequently from calypterate 
Diptera breeding in such habitats as fungus, 
dung, flower heads and other seed bearing 
structures. A few have also been recorded 
from acalypterates such as drosophilids and 
sciomyzids (Papp 1972, Fischer 1975, 
Wharton 1984, Vet and van Alphen 1985, 
van Achterberg 1988). Hosts have not been 
previously recorded for any of the Neotrop- 
ical species of Phaenocarpa. 

Materials and Methods 

Terminology is as in Wharton (1980, 
1994) except as follows: venation conforms 



198 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



to the Standards recently established by 
Sharkey and Wharton (1997); scutellar sul- 
cus is used in place of prescutellar pits; the 
terms mesonotal disc (or simply disc) and 
anterior declivity are retained in their for- 
mer meaning, but are referred to as parts of 
the mesoscutum. Measurements are given 
to the nearest 0.05. 

Museum acronyms are as follows: The 
Natural History Museum, London 
(BMNH), Texas A&M University Insect 
Collection (TAMU). Part of the material for 
this study was sorted from general BMNH 
accessions by RAW, the remainder was 
reared by NSC as part of a program to de- 
termine natural enemies of fruit-infesting 
tephritid flies of the genus Anastrepha in 
Colombia. 

Key TO Neotropical Species of 
Phaenocarpa 

1. Second fiagellomere at least 1.4 times longer 
than first (Figs. 12-14) 2 

- Second fiagellomere at most 1.1 times longer 
than first (Fig. 1 ) 6 

2. Second submarginal cell short, with 2RS near- 
ly equal to or slightly longer than 3RSa .... 3 

- Second submarginal cell longer (Fig. 7), with 
3RSa at least 1.25 times longer than 2RS ... 4 

3. Notauli distinct posteriorly. Mandible with 
tooth I extending distinctly distad tooth 3 . . . 

Phaenocarpa co.xalis (Szepligeti) 

- Notauli absent posteriorly, not reaching midpit 
on mesoscutum. Mandible with tooth 3 extend- 
ing distinctly distad tooth I 

Phaenocarpa anomala Wharton 

4. Mandible with broad tooth 1 separated from 
tooth 2 by a deep cleft (Fig. 15). Fore wing 
(RS + M)b distinctly more than half length of 
m-cu (Fig. 7) 

Phaenocaipa pericarpa Wharton and Carrejo, n. sp. 

- Mandibular teeth 1 and 2 connected by a 
broad, undulant flange, not separated from each 
other by a deep cleft (Fig. 10). Fore wing 
(RS + M)b short to absent, distinctly shorter 
than half length of m-cu 5 

5. Petiole dark brown, narrowly elongate, about 

1.5 times longer than apical width; apex about 

1.6 times wider than base. Propodeal areola 
distinct; transverse carina of propodeum absent 
laterally Phaenocarpa heynei Papp 

- Petiole dark yellow, broader, about 1.2 times 
longer than apical width, apex about twice wid- 
er than base. Propodeal areola indistinct; trans- 



verse carina complete to spiracles 

Phaenocarpa siibtilistriata Papp 

6. Stigma of fore wing broad distally, abruptly 
narrowing basad junction with r in females 
(Fig. 5), with posterior margin indistinct on 
basal half; divided by a hyaline, desclerotized 
line in males, with portion of stigma posteriad 
hyaline region broader than portion along an- 
terior margin of wing (Fig. 6). Mesopleuron, 
metapleuron, and propodeum dark brown. 
Metanotum in lateral view with short but dis- 
tinct spine (Fig. 2) 

Phaenocarpa hyalina Trostle. n. sp. 

- Stigma of fore wing narrow distally and basal- 
ly, only gradually narrowing basad junction 
with r in females, with posterior margin clearly 
delineated on basal half; males in which the 
stigma is divided by a desclerotized line have 
the portion that is posteriorad the desclerotized, 
hyaline region narrower than the portion along 
the anterior margin of the wing. Body color 
and development of metanotal projection var- 
iable 7 

7. Mesopleuron, metapleuron, and propodeum 
yellow. Metanotum in lateral view with short, 
narrow spine-like projection (as in Fig. 2) . . . 

Phaenocarpa sharkeyi Wharton 

- Mesopleuron, metapleuron, and propodeum 
dark brown, contrasting with light brown to 
yellow mesoscutum. Metanotum in lateral view 
with broadly triangular projection (distinctly 
broader than high) 

Phaenocarpa cratomorpha Wharton 



Phaenocarpa hyalina Trostle, 
new species 

(Figs. 1-6) 

Female. — Head: Moderately transverse 
in dorsal view, 1.3X wider than long. Face 
punctate, setiferous but polished, 1.25-1.35 
higher than wide. Frontoclypeal suture 
broad, relatively shallow, unsculptured. 
Clypeus broad, weakly convex, barely pro- 
truding. Frons smooth, bare, nearly flat, 
weakly concave medially. Mandible 1.85X 
longer than width between tooth 1 and 3, 
surface largely smooth; tooth 1 broad, near- 
ly orthogonal, with dorsal margin concave; 
distinct cleft present between tooth 1 and 2, 
tooth 2 lacking additional tooth or knob; 
tooth 2 acutely triangular, short, 0.4-0.45 X 
apical width of mandible, but extending be- 
yond tooth 1 and 3; tooth 3 broadly than- 



VOLUME 101, NUMBER I 



199 




Figs. 1-4. Phaenocarpa hyalina. 1. Antenna showing relative lengths of basal 3 flagellomeres. 2, Mesosoma, 
lateral view, showing metanotal spine. 3, Mesosoma, dorsal view. 4. Metasoma, dorsal view. 



gular. Eye large, 4.25-5 X longer than tem- 
ple. Anterior tentorial pit large, extending 
more than 0.5 X distance from lateral mar- 
gin of clypeus to eye. Antenna 25-27 seg- 
mented; second flagellomere 1.0-1. 05 X 



longer than first; first flagellomere 1.1-1.3X 
longer than third. Eye-antennal sulcus ab- 
sent. 

Mesosoma: 1.35-1. 5 X longer than high. 
Pronotum predominately smooth, with Ion- 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





8 



Figs. 5-8. 
wing. 



Wings. 5, Phaenocarpu hycilina. female. 6, P. hyciliiui. male. 7, 8, P. pericarpa fore and hind 



gitudinal carina separating dorsal and lat- 
eral portions, and fine crenulae laterally ex- 
tending ventrally from this carina; prono- 
tum dorsally a narrow band with a shallow, 
barely perceptible median pit; anterior mar- 
gin weakly emarginate. Mesoscutum pol- 



ished; anterior declivity vertical, sparsely 
setose; notauli extending medially as cari- 
nate ridges along dorsal margin of anterior 
declivity, then proceeding posteriorly as 
weak, parallel grooves perpendicular to the 
carinate ridges, notaular grooves evanescent 



VOLUME 101, NUMBER 1 



201 



near anterior margin of midpit; 3-4 pairs of 
setae extending from transverse portion of 
notauli to transscutal articulation; midpit 
well developed, extending anteriorly 0.4 X 
length of disc from transscutal articulation. 
Scutellar sulcus about 2X broader than 
long, with medial carina; lateral margins of 
axilla flangelike. Metanotum with posterior 
margin a broad, flat plate, elevated medially 
as a prominent spine, median field with or 
without carinae radiating from spine. Pro- 
podeum areolate, as in other members of 
the cratomorpha species group. Mesopleu- 
ron with small patch of rugose sculpture on 
anterior margin medially, otherwise 
smooth, polished, virtually bare; carinately 
margined posteriorly. Sternaulus sinuate, 
crenulate over anterior 0.65, smooth over 
posterior 0.35. Metapleuron largely un- 
sculptured, as in P. sharkeyi. 

Wings: Fore wing stigma 4. 5-5. OX lon- 
ger than broad, solid throughout (i.e., with- 
out hyaline streak), basal half distinctly nar- 
rower than distal half with posterior margin 
of basal half indistinct and somewhat ex- 
cavated, stigma abruptly widening near 
junction with r; r short, slightly shorter than 
and arising slightly distad mid-width of 
stigma; 3RSa 1.4-1. 8X longer than 2RS; 
3RSb extending to wing tip, weakly curved 
at apex; (RS + M)b usually absent; Icu-a of- 
ten very short, postfurcal by 1.25-3. 5 X its 
length. Hind wing with IM 2.4-3.5 X lon- 
ger than M + CU; 3 hamuli. 

Metasoma: Petiole 1.6-1.7X longer than 
apical width, apex 1.5-1.6X v/ider than 
base, dorsal carinae on petiole more or less 
parallel sided, weakly converging and be- 
coming less prominent posteriorly; surface 
sculpture often granular- rugose medially; 
more finely rugose laterally. Ovipositor 
sheath with 3 irregular rows of approxi- 
mately 7 setae each. Ovipositor 0.9 X and 
ovipositer sheath 0.65 X length of mesoso- 
ma. 

Color: Generally brown; mesoscutum, 
metanotum and occasionally propleuron 
bright orange yellow; mesopleuron, meta- 
pleuron, propodeum and petiole dark 



brown; legs white at coxae and gradually 
darkening to yellow distally; maxillary palp 
white; face variable, dark brown to light or- 
ange yellow, usually paler near antennae; 
scape and pedicel bright yellow with pedi- 
cel slightly paler; basal 16-18 flagellomeres 
gradually darkening from yellow to brown 
distally, followed by 4-7 white flagello- 
meres, and terminating with 1-3 (usually 
1.5) brown flagellomeres; wings hyaline. 

Body length: 2.1-2.4 mm 

Male. — Similar to female except as fol- 
lows: head 1.3-1.4X wider than long; face 
1.1-1.2X higher than wide; mesosoma 
1.45-1. 55 X higher than wide; body 2.3-2.9 
mm; fore wing stigma longitudinally bi- 
sected for most of its length with the thick- 
ened portion along anterior margin of wing 
almost completely separated from posterior 
portion by hyaline streak, the two portions 
only narrowly joined distally, posterior por- 
tion broader than anterior portion. 

Biology. — Unknown 

Material examined. — Holotype $ : 
"Brasilien Nova Teutonia 27°irB. 
52°23'L. 2-ix-1935 Fritz Plaumann" 
(BMNH). Paratypes (BMNH, TAMU): 2 S, 
same data as holotype; 2 $, 10 6, same 
except viii.1935, xi.l935, 6.iii.l936, and 
15.x. 1935; 1 9,2 6, same locality but 
country spelled "Brazil:" and with B.M 
Accession label "B.M. 1937-656", collect- 
ed ll.vii.l937, and 7.V.1937; 2 $, 2 d, 
same as preceding, but "B.M. 1938-682", 
collected 20.iv.l938, 13.viii.l938, 
23.iv.1938, and 3.V.1938. 

Diagnosis. — This species is a member of 
the cratomorpha species group as defined 
by Wharton (1994). As in other species of 
this group, P. hyalina has the second fla- 
gellomere equal to or barely longer than the 
first (Fig. 1), a median metanotal projection 
(Fig. 2), and a hyaline streak longitudinally 
bisecting the male stigma (Fig. 6). The stig- 
ma is sexually dimorphic, lacking a well- 
defined streak in females. Phaenocarpa hy- 
alina differs from other described species 
of this group in the possession of a broader 
stigma that narrows more abruptly basad 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



the junction with r. Further, the metanotal 
projection of P. hyalina is much more spi- 
nose than in P. cratomorpha, displaying a 
thinner, finer apex. Obvious color differenc- 
es exist between P. sharkeyi and P. hyalina. 
Phaenocarpa sharkeyi is more uniformly 
pale yellow in coloration, contrasting with 
the darker brown portions of the head, 
meso- and metasoma of P. hyalina. 

Discussion. — This species is named for 
the hyaline streak that almost completely 
bisects the stigma in males, resulting in sep- 
arate anterior and posterior portions. Addi- 
tional differences in size and shape between 
males and females were noted by Wharton 
(1994) in his description of P. cratomor- 
pha. Detailed scrutiny of males and females 
of P. hyalina, however, fails to reveal sig- 
nificant differences between the sexes for 
most of these same characters. Since so few 
individuals of P. cratomorpha were avail- 
able for study, it is quite possible that the 
differences recorded by Wharton (1994) in 
the size and shape of the eye and petiole 
were simply intraspecific rather than spe- 
cifically intersexual. Measurements of in- 
dividuals from the much longer series of P. 
hyalina demonstrate that these quantitative 
characters are highly variable within each 
sex. 

Although a distinct hyaline streak is pre- 
sent in males and absent in females, the 
stigma in females of P. hyalina exhibits 
modifications suggesting a morphocline 
leading to the extreme condition expressed 
in males. The stigma in females has a slight 
indication of desclerotization in a position 
comparable to the basal portion of the hy- 
aline streak in males. Further, the stigma is 
exceptionally narrow basally, distad the 
parastigma, then somewhat abruptly widens 
where it meets r. In most other species of 
Phaenocarpa, the posterior margin of the 
stigma is straight to evenly convex between 
r and the parastigma. Phaenocarpa hyalina 
and the species described below are excep- 
tional in this regard. 



Phaenocarpa pericarpa Wharton and 
Carrejo, new species 

(Figs. 7, 8, 11, 14-16, 19) 

Female. — Head: Moderately transverse 
in dorsal view, about 1.5-1.6X wider than 
long; eyes strongly bulging beyond tein- 
ples. Face finely punctate, the punctations 
separated by much more than their diame- 
ter, otherwise smooth and polished between 
base of antenna and frontoclypeal suture; 
frontoclypeal suture deep and very wide, 
transversely striate; clypeus exceptionally 
narrow and strongly protruding, lateral mar- 
gin of clypeus distinctly separated from an- 
terior tentorial pit, the latter small, set with- 
in broad concavity between clypeus and 
lower margin of eye, the concavity trans- 
versely striate at least in part. Malar space 
absent. Frons smooth, polished, evenly and 
weakly convex, bare except for 2-4 short 
setae along eye margin laterad ocelli. Ver- 
tex bare. Mandible 1.4-1.5X longer than 
apical width, distinctly expanded apically, 
apex 1.7-1.85X wider than base, surface 
with strigose sculpture extending from mid- 
dle of diagonal ridge to dorsal margin at 
base of tooth 1 ; tooth 1 very broad, orthog- 
onal, separated by deep cleft from narrowly 
triangular tooth 2; tooth 2 without dorsal 
knob; tooth 3 extending distally not quite 
as far as tooth 1 , narrower and more round- 
ed than the latter. Eye very large, 5. 2-6.0 X 
longer than temple, with a few, scattered, 
minute setae visible at 50 X. Antenna 39 
segmented; second flagellomere 1.5-1.7X 
longer than first, 1 . 1 X longer than third. 
Maxillary palp very long, about twice 
height of head. 

Mesosoma: 1. 45-1. 55 X longer than 
high. Pronotum dorsally smooth, polished, 
but uneven: with small, u-shaped depres- 
sion medially along anterior margin; weak- 
ly elevated as a small, rounded bump pos- 
teromedially; flattened laterally. Mesoscu- 
tum smooth, polished, with setae confined 
to a short row along notauli on anterior de- 
clivity, absent on disc; notauli shallow, 
weakly sculptured, confined to anterior de- 



VOLUME 101. NUMBER 1 



203 




Figs. 9-16. Phaenocarpa spp. 9-11, Faces. 9. P. siibtilistriata. 10. P. heyuei. 11. P. pericarpa. 12-14, 
Antennae showing relative lengths of basal 3 flagellomeres (setal pattern shown only on Fig. 13). 12. P. subtil - 
isrriatci. 13, P. Iieynei. 14, P. pericarpa. 15, P. pericarpa. mandible. 16, P. pericarpa, propodeum. 



clivity; midpit small, shallow, oval. Scutel- 
lar sulcus twice as wide as long, or nearly 
so; with a single median ridge; lateral fields 
polished, unsculptured. Metanotum with 
thin, longitudinal flange along midline, the 
flange sloping more precipitously anteriorly 
than posteriorly, not elevated above level of 
scutellum. Propodeum smooth, polished, 
with well-defined pentagonal areola delim- 
ited by strong carinae; areola narrow, about 
half as wide as tall, confined to posterior 
half of propodeum; anterior half with a 
strong median carina; lateral carina extend- 
ing from areola to spiracle usually very 
weak over lateral half. Sternaulus long, nar- 
row, sinuate, complete from anterior margin 
to mid coxa, weaker posteriorly; crenulate 



anteriorly, the sculpture weakening poste- 
riorly and usually absent over posterior 0.3- 
0.5. Metapleuron finely punctate but oth- 
erwise polished and unsculptured over most 
of surface. 

Wings: Fore wing stigma weakly con- 
cave basally along posterior margin, grad- 
ually widening distally towards junction 
with r, solid throughout, about 5X longer 
than width at r, r arising from distal 0.7- 
0.75; r very short, 0.1 5-0.20 X length of 
2RS; 2RS sharply angled near posterior 0.2; 
3RSa 1.25-1. 35 X longer than 2RS, second 
submarginal cell gradually narrowing dis- 
tally; 3RSb ending at wing tip, weakly and 
evenly bowed; m-cu distinctly antefurcal, 
with (RS + M)b 0.65-0.8 X length of m-cu; 



204 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Icu-a postfurcal by 2-3 X its length; 2CU 
interstitial: arising directly in line with 
ICU. Hind wing very narrow, about 6X 
longer than wide; IM 2.0-2.5 X longer than 
M+CU; 2M very short, angled towards 
posterior margin, m-cu absent. 

Metasoma: Petiole 1.3-1.45X longer 
than apical width; apex 1.8-1.9X wider 
than base; surface strigose, the sculpture 
distinct medially, often weak laterally; dor- 
sal carina strong basally, evanescent at level 
of spiracles, absent posteriorly; dorsope 
large and deep. Remaining terga without 
sculpture. Ovipositor of moderate length, 
ovipositor sheath not fully exposed in ma- 
terial available for examination, but about 
2.0-2.3 X longer than mesosoma; ovipositor 
finely tapered to apex, without discernible 
subapical node or notch. 

Color: Dark brown; mandible, scape, 
pedicel, propleuron and petiole variously 
lighter brown or red-brown in most speci- 
mens; flagellomeres 1-4 brown, 5-12 and 
19-29 dark brown. 14-18 white, and 13 bi- 
colored brown and dark brown; fore and 
mid legs and hind coxa and trochanter yel- 
low to dark yellow, hind femur mostly yel- 
low with dark spot dorsally over apical 0.3- 
0.4, hind tibia and tarsus brown; hypopy- 
gium apically and apical tergite yellow or 
yellow brown; palps white. 

Male. — A single male, probably repre- 
senting this species, fits the above descrip- 
tion except as follows: face shorter, about 
1.5X wider than high; mandible less ex- 
panded distally, about 1.8X longer than api- 
cal width; fore wing Icu-a postfurcal by 
only about 1.4X its length; and fore wing 
3RSa about 1.6X 2RS. Flagellomeres 15 
and 16 are dirty white, with remaining fla- 
gellomeres dark brown. 

Biology. — Reared from puparia of An- 
astrepha distincto Greene in pods of guama 
{Inga sp.: Fabaceae). 

Material examined. — Holotype 9 : "CO- 
LOMBIA Dept. del Valle del Cauca Mun. 
Buenaventura, Corregimiento de Zacarias 
23.iii.1994 N. Carrejo ex Anastrepha dis- 
tincta on Guama" Deposited in La Univer- 



sidad del Valle, Museo de Entomologia. 
Paratopes: 3 9 , same data as holotype 
(TAMU; Universidad del Valle; and Insti- 
tuto de Ciencias Naturales, Universidad 
Nacional de Colombia). Additional material 
(not a paratope): 1 6, VENEZUELA, Ar- 
agua, Parque Nacional Henri Pittier, Ran- 
cho Grande, 1,100m, 12.1.1996, R. Wharton 
(TAMU). 

Diagnosis. — As in both P. heynei and P. 
subtilistriata, this species has reduced no- 
tauli, a short first flagellomere, and a large 
second submarginal cell. It differs from 
both of these species by the possession of 
a shorter ovipositor (compare Fig. 19 with 
Figs. 17, 18) which lacks a subapical node 
or notch and a broader, more discrete tooth 
1 on the mandible (Fig. 15). The ovipositor 
is more than three times longer than the me- 
sosoma in P. heynei and P. subtilistriata. 
The shape of the mandible is sufficient for 
separating P. pericarpa from all other de- 
scribed New World species of Phaenocar- 
pa. 

Discussion. — Papp (1969) adequately 
differentiated P. heynei from P. subtilistria- 
ta, noting especially the difference in shape 
of the petiole. There are also slight differ- 
ences in sculpture between the holotypes of 
P. heynei and P. subtilistriata. The propo- 
deal areola is essentially obliterated in P. 
subtilistriata, with the posterior face stri- 
gose or weakly rugulose below the well- 
developed and complete transverse carina. 
A weak areola is present in P. heynei, and 
the transverse carina is incomplete, not 
reaching the propodeal spiracle. The ster- 
naulus is also broader and more heavily 
sculptured in P. subtilistriata but the scu- 
tellar sulcus is smooth with a single median 
carina. In P. heynei, the scutellar sulcus is 
weakly sculptured on either side of the me- 
dian sulcus. Although both P. heynei and 
P. subtilistriata are known only from the 
holotype, and variation thus cannot be as- 
sessed, these same sculptural features show 
little vaiiation in the four specimens of P. 
pericarpa. Thus, it is likely that these rel- 
atively minor sculptural differences will be 



VOLUME 101. NUMBER 



205 




Figs. 17-19. Mcso- and melasoma, lateral view. 17, Phaenuccupa suhlilistriata. 18, P. heyiwi. 19, P. pcri- 
carpa. 



useful for species-level recognition of P. 
heynei and P. subtilistriata. 

Several features suggest that P. heynei 
shares a sister group relationship with P. 
pericarpa relative to P. subtilistriata. Al- 
though the clypeus is narrow in all three 
species, it is more strongly narrowed and 
protruding in P. heynei and P. pericarpa 
(Figs. 10, 11), with consequent detachment 
of the anterior tentorial pit from the lateral 
margin of the clypeus. Similarly, 2M in the 
hind wing is shorter and more distinctly de- 
flected posteriorly in P. heynei and P. peri- 
carpa than in P. subtilistriata, which retains 



the more plesiomorphic form of a longer, 
more distally-directed 2M. The fore wing 
stigma of P. heynei is also more similar to 
the unusually shaped stigma off*, pericarpa 
than the more typically shaped stigma of P. 
subtilistriata. Phaenocarpa subtilistriata 
appears to form a link between the hey- 
nei^ pericarpa sister group and the more 
typical Phaenocarpa species from the Hol- 
arctic Region. This hypothesized relation- 
ship between P. heynei and P. pericarpa 
leads the suggestion that the Gnathopleura- 
like mandible of P. heynei and P. subtilis- 
triata is a more primitive feature from 



206 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



which the P. pericarpa configuration was 
derived. Despite similarities in the mandi- 
ble, P. heynei and P. subtilisthata can be 
readily separated from Gnathopleiira since 
the latter has a much smaller second sub- 
marginal cell. 

The putative relationship between P. hey- 
nei and P. pericarpa is based on features 
which, though unusual, are found elsewhere 
in Phaenocarpa and related genera. A sim- 
ilarly narrow hind wing with reduced 2M 
is found in the Phaenocarpa cratomorpha 
species group, for example. A narrow, 
strongly protruding clypeus is found in at 
least two species oi Asobara (one from Bra- 
zil and one from Papua New Guinea) which 
otherwise lack shared derived features. In 
all of these cases, the character states in 
question are hypothesized as independent 
derivations. 

This is the first host record for any of the 
Neotropical species of Phaenocarpa. The 
type series of P. pericarpa was reared from 
the tephritid fly Anastrepha distincta 
Greene, developing in the pods of guama 
{Inga sp.). Relatively few tephritids have 
been recorded as hosts of Alysiinae (Whar- 
ton 1984). Only one, Asobara anastrephae 
(Muesebeck), has been verified as a para- 
sitoid of fruit-infesting tephritids. Asobara 
orientalis Viereck, originally described 
from material thought to be reared from da- 
cine tephritids, is undoubtedly a drosophilid 
parasitoid. Three other alysiine species have 
been reared either from tephritids in flower 
heads or other plant parts, and at least four 
additional species have been reared from 
unknown hosts in fruit. Asobara anastre- 
phae is interesting because of its exception- 
ally large size and unusual host preferences 
relative to other species of Asobara, which 
are primarily drosophilid parasitoids. Aso- 
bara anastrephae belongs to a group of 
Neotropical species with typical Asobara 
fore wing venation, loss of hind wing cu-a, 
a short, broad petiole, and brightly colored 
bodies (Muesebeck 1958, Wharton 1994). 
One of the derived members of this group, 
as yet undescribed, exhibits the same clyp- 



eal modifications found in P. pericarpa and 
P. heynei. 

Acknowledgments 

We are grateful to Tom Huddleston (for- 
merly BMNH) for extending many cour- 
tesies regarding material in his care, to 
Frank Koch (Humboldt Museum, Berlin), 
Jeno Papp (Hungarian Natural History 
Museum), and Paul Marsh and David 
Smith (Systematic Entomology Laborato- 
ry, USDA) for the loan of type material 
and assistance with type collections. Barry 
Flahey provided the illustrations for P. hy- 
alina. This work was supported in part by 
the Texas Agricultural Experiment Station 
and by the National Science Foundation 
(DEB 9300517). 

Literature Cited 

Achterberg, C. van. 1988. The genera of the Aspilota- 
group and some descriptions of fungicolous Aly- 
siini from the Netherlands (Hymenoptera: Bracon- 
idae: Alysiinae). Zoologische Verhandelingen 
247: 1-88. 

Fischer, M. 1971. Untersuchungen iiber die Euro- 
paischen Alysiini mit besonderer Berucksichti- 
gung der Fauna Niederosterreichs. Polskie Pismo 
Entomologiczne 41: 19-160. 

. 1974. Die nearktischen Phaenocarpa- Arten. 

Revision der Gruppe B (Hymenoptera, Braconi- 
dae, Alysiinae). Polskie Pismo Entomologiczne 
44: 103-230. 

. 1975. Die nearktischen Phaenocarpa- Arlen. 

Revision der Gruppe A (Hymenoptera. Braconi- 
dae, Alysiinae). Polskie Pismo Entomologiczne 
45: 279-356. 

. 1990. Westpalaarktische Phaenocarpa- Arten: 

Vorlaufiger Bestimmungsschlussel, Deskriptionen 
und Redeskriptionen (Hymenoptera, Braconidae, 
Alysiinae). Annalen Naturhistorische Museum in 
Wien 91: 105-135. 

. 1993. Einige Phaenocarpa-Wespen aus der 

Alten Welt: Redeskriptionen und Stellung in ei- 
nem vergleichenden System (Hymenoptera. Bra- 
conidae, Alysiinae). Linzer biologische Beitrage 
25: 511-563. 

. 1994. Beitrag zur Kenntnis der Keiferwespen 



der Welt (Hymenoptera, Braconidae, Alysiinae: 
Alysiini). Linzer biologische Beitrage 26: 763- 
806. 
Muesebeck, C. F. W. 1958. New Neotropical wasps of 
the family Braconidae (Hymenoptera) in the U.S. 
National Museum. Proceedings of the United 
States National Museum 107: 405-461. 



VOLUME 101. NUMBER 1 



207 



Papp, J. 1966. New Phaenocarpa Forster species from 
the Ethiopian Region (Hymenoptera, Braconidae). 
Acta Zoologica Academiae Scientiarum Hungari- 
cae 12: 133-144. 

. 1969. A synopsis of the Phaenocarpa Forst. 

species of the Neotropic Region (Hymenoptera: 
Braconidae, Alysiinae). Acta Zoologica Acade- 
miae Scientiarum Hungaricae 15: 379-389. 

. 1972. Phaenocarpa impugnata sp. n. (Hym., 



Braconidae: Alysiinae), a new reared species from 
Denmark. Zoologischer Anzeiger, Leipzig 188: 
52-56. 

Sharkey, M. J. and R. A. Wharton. 1997. Morphology 
and terminology, pp. 19-63. //; Wharton, R. A., 
P. M. Marsh and M. J. Sharkey, eds.. Manual of 
the New World genera of the family Braconidae 
(Hymenoptera). Special Publication No. 1 of the 
International Society of Hymenopterists. 

Shenefelt. R. D. 1974. Pars 11. Braconidae 7 Alysi- 
inae, pp. 937-1113. In van der Vecht, J. and R. 
D. Shenefelt, eds., Hymenopterorum Catalogus 
(nova editio). Dr. W. Junk, The Hague. 



Tobias, V. L 1986. Subfamily Alysiinae, pp. 100-231. 
In Identification of Insects of European USSR, 
Vol. Ill, Part V, Hymenoptera, Braconidae. Aka- 
demia Nauk, Leningrad (in Russian). 

Vet, L. E. M. and J. J. M. van Alphen. 1985. A com- 
parative functional approach to the host detection 
behaviour of parasitic wasps. 1. A qualitative 
study on Eucoilidae and Alysiinae. Oikos 44: 
478-486. 

Wharton, R. A. 1980. Review of the Nearctic Alysiini 
(Hymenoptera, Braconidae) with discussion of ge- 
neric relationships within the tribe. University of 
California Publications in Entomology 88: 1-112. 

. 1984. Biology of the Alysiini (Hymenoptera: 

Braconidae), parasitoids of cyclorrhaphous Dip- 
tera. Texas Agricultural Experiment Station Tech- 
nical Monograph 11: 1-39. 

. 1994. New genera, species, and records of 

New World Alysiinae (Hymenoptera: Braconi- 
dae). Proceedings of the Entomological Society of 
Washington 96: 630-664. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999. pp. 208-211 

THE SMALL MINNOW MAYFLY GENUS CLOEODES TRAVER 
(EPHEMEROPTERA: BAETIDAE) IN MADAGASCAR 

C. R. Lugo-Ortiz, W. P. McCafferty, and J.-L. Gattolliat 

(CRL, WPM) Department of Entomology, Purdue University, West Lafayette, IN 
47907, U.S.A. (e-mail: carlosJugo-ortiz@entm.purdue.edu); (JLG) Museum of Zoology, 
PO. Box 448, CH 1000 Lausanne, Switzerland 



Abstract. — Cloeodes portabilis, new species, represents the first report of Cloeodes 
from Madagascar The species is distinguished in the larval stage by the relatively wide 
anteromedial emargination of the labrum, presence of tufts of fine, simple setae between 
the prosthecae and molae of the mandibles, slightly distolaterally acute segment 3 of the 
labial palps, and abdominal color pattern. The presence of Cloeodes throughout the South- 
em Hemisphere suggests a relatively ancient origin among extant Baetidae. 

Key Words: Ephemeroptera, Baetidae, Cloeodes portabilis, new species, Madagascar 



The distinctive small minnow mayfly ge- 
nus Cloeodes Traver (Ephemeroptera: Bae- 
tidae) has been known from the Afrotrop- 
ics, Neotropics, Orient, and southwestern 
Nearctic (Traver 1938, Waltz and Mc- 
Cafferty 1987ab, 1994, Kluge 1991, Flow- 
ers 1991, Lugo-Ortiz and McCafferty 1993, 

1994, 1995, McCafferty and Lugo-Ortiz 

1995, McCafferty et al. 1997). Most re- 
cently, Lugo-Ortiz and McCafferty (1998) 
reported Cloeodes from Australia, signifi- 
cantly extending its known range. Three 
historic biogeographic hypotheses were 
provided to explain its essentially Pantrop- 
ical distribution. 

That Cloeodes was well established 
throughout Gondwanaland or at least West 
Gondwanaland (the South American- Afri- 
can-Malagasy-Indian landmass) prior to 
100 million years ago, before Madagascar 
began to separate from Africa and India, 
were hypotheses that clearly predicted that 
Cloeodes would occur in Madagascar. 
Herein we report Cloeodes from Madagas- 
car for the first time based on a new species 
described from larvae. An additional spe- 



cies of Cloeodes from the island will be de- 
scribed elsewhere by one of us (J.-L.G.). 
The specimens studied are housed in the 
Purdue Entomological Research Collection, 
West Lafayette, Indiana. 

Cloeodes portabilis Lugo-Ortiz and 
McCafferty, new species 

(Figs. 1-10) 

Larva. — Body length: ?)A-AA mm. Cau- 
dal filaments length: 1.8-2.0 mm. Head: 
Coloration medium brown, with no distinct 
pattern. Antenna approximately 1.75X 
length of head capsule. Labrum (Fig. 1) 
with wide anteromedial emargination, sub- 
medial pair of long, fine, simple setae, and 
submarginal row of four to six long, fine, 
simple setae. [Left and right mandibles 
(Figs. 2, 3) with incisors worn down.] Left 
mandible (Fig. 2) with one set of incisors; 
prostheca robust, apically denticulate; tuft 
of fine, simple setae present between pros- 
theca and mola. Right mandible (Fig. 3) 
with two sets of incisors; prostheca slender, 
apically bifid; tuft of fine, simple setae pre- 
sent between prostheca and mola. Maxilla 



VOLUME 101. NUMBER 



209 







j/umMi^ 






WAA/'^^^^^^'^^V\A/\AAAAAA/ 



8 





10 



Figs. 1-10. Cloeodes portabilis. 1, Labrum (dorsal). 2, Left mandible. 3, Right mandible. 4, Left maxilla. 
5, Labium (left-ventral; right-dorsal). 6, Left foreleg. 7, Abdomen (dorsal). 8, Tergum 3 (detail). 9, Gill 3. 10, 
Paraproct. 



(Fig. 4) with four small, stout denticles on 
crown of galealacinia; four to five long, 
fine, simple setae near medial hump; palp 
not reaching galealacinia; palp segment 1 
subequal in length to segment 2; segment 2 
apically acute. Labium (Fig. 5) with glossa 



and paraglossa equal in length; palp seg- 
ment 1 approximately 0.80 X length of seg- 
ments 2 and 3 combined; segment 2 ap- 
proximately 1.6X length of segment 3; seg- 
ment 3 falcate apically and slightly pointed 
distolaterally. Thorax: Coloration yellow 



210 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



brown, with complex markings. Hindwing- 
pads absent. Legs (Fig. 6) pale yellow 
brown; femora dorsally with row of five to 
six medium-sized, robust, simple setae in- 
termixed with numerous long, fine, simple 
setae; tibiae ventrally with 10-12 short, 
stout, simple setae; tarsi ventrally with eight 
to nine short, stout, simple setae. Abdomen 
(Fig. 7): Coloration medium to yellow 
brown; segment 1 yellow brown; segment 
2 yellow brown, with submedial medium 
brown subtriangular marking and oblique 
medium brown distolateral markings; seg- 
ment 3 with submedial medium brown cres- 
centlike marking posteriorly and oblique 
medium brown distolateral markings; seg- 
ment 4 with submedial medium brown cres- 
centlike marking posteriorly; segment 5 
with large, wavy, medium brown marking 
posteriorly; segment 6 with submedial 
spikelike medium brown marking and 
oblique medium brown distolateral mark- 
ings; segment 7 with small, oblong, medi- 
um brown anterolateral markings; segment 
8 yellow brown, with no markings; seg- 
ments 9-10 medium brown, with no mark- 
ings. Terga (Fig. 8) with abundant scale ba- 
ses, and with posterior marginal spines tri- 
angular, approximately as long as basal 
width. Sterna yellow brown. Gill (Fig. 9) 
subtriangular, well tracheated, marginally 
smooth. Paraproct (Fig. 10) with 10-12 
spines, increasing in size apically. Caudal 
filaments pale yellow brown; medial caudal 
filament subequal in length to cerci. 

Adult. — Unknown. 

Material examined. — Holotype: Larva, 
MADAGASCAR, Tamatave (= Toamasi- 
na) Prov., stream at Gri-Gri, RN 2, 17-X- 
1971, G. E, C. H. Edmunds, and E Em- 
manuel. Paratopes: Two larvae, same data 
as holotype; five larvae, MADAGASCAR, 
Antsiranana Prov., Djabala R., 1 1 km NW 
of Hell-Ville, Nosy Be, 25-X-1971, G. E, 
C. H. Edmunds, and F Emmanuel [mouth- 
parts, left foreleg, tergum 3, gills 3, and 
paraproct of one larva mounted on slide 
(medium: Euparal)]. Additional material: 
Four larvae, same data as holotype; twenty 



larvae, MADAGASCAR, Antsiranana 
Prov., Djabala R., 1 1 km NW of Hell-Ville, 
Nosy Be, 25-X-1971, G. E, C. H. Edmunds, 
and E Emmanuel. 

Etymology. — The specific epithet is a 
Latin word meaning "that which may be 
carried." It is an allusion to the species be- 
ing a drifted representative of the genus. 

Discussion. — Cloeodes portabilis is dis- 
tinguished from other members of the ge- 
nus by the relatively wide anteromedial 
emargination of the labrum (Fig. 1), pres- 
ence of tufts of fine, simple setae between 
the prosthecae and molae of the mandibles 
(Figs. 2, 3), slightly distolaterally acute seg- 
ment 3 of the labial palps (Fig. 5), and ab- 
dominal color pattern (Fig. 7). 

Cloeodes portabilis is unique among 
Eastern Hemisphere members of the genus 
because segment 3 of the labial palps is 
somewhat falcate, being slightly distolater- 
ally pointed (Fig. 5); other species have a 
bulbous segment 3. Only the South Amer- 
ican species C. hydation McCafferty and 
Lugo-Ortiz has a similar labial palp mor- 
phology (McCafferty and Lugo-Ortiz 1995: 
Fig. 6); we can only assume that it consti- 
tutes a homoplasy. In addition, the presence 
of a tuft of setae between the prosthecae 
and molae of the mandibles in C. portabilis 
(Figs. 2, 3) is exceptional in Cloeodes, pos- 
sibly indicating a relatively ancestral posi- 
tion within the genus, because that charac- 
teristic is generally associated with plesi- 
otypic genera in Baetidae (R. D. Waltz, per- 
sonal communication). 

Acknowledgments 

We thank G. E Edmunds, Jr., Salt Lake 
City, Utah, for the donation of the material 
used in this study. This paper has been as- 
signed Purdue Agricultural Research Pro- 
gram Journal No. 15655. 

Literature Cited 

Flowers, R. W. 1991. Diversity of stream-living in- 
sects in northwestern Panama. Journal of the 
North American Benthological Society 10: 

322-334. 



VOLUME 101. NUMBER 1 



211 



Kluge, N. 1991. Cuban mayflies of the family Bae- 
tidae (Ephemeroptera) 1. Genera Collibaetis. 
Cloeodes, and Paracloeodes. Zoologischeskiy 
Zhurnal 12: 128-136. [in Russian] 

Lugo-Ortiz. C. R. and W. R McCafferty. 1993. Gen- 
era of Baetidae (Ephemeroptera) from Central 
America. Entomological News 104: 193-197. 

. 1994. New records of Ephemeroptera from 

Mexico. Entomological News 105: 17-26. 

. 1995. Annotated inventory of the mayflies 

(Ephemeroptera) of Arizona. Entomological 
News 106: 131-140. 

. 1998. First report and new species of the 



genus Cloeodes (Ephemeroptera: Baetidae) 
from Australia. Entomological News 109: 122- 
128. 
McCafferty, W. R and C. R. Lugo-Ortiz. 1995. 
Cloeodes hydation. n. sp. (Ephemeroptera: Bae- 
tidae): an extraordinary, drought tolerant may- 



fly from Brazil. Entomological News 106: 29- 
35. 

McCafferty. W. R, C. R. Lugo-Ortiz, and G. Z. Ja- 
cobi. 1997. The mayfly fauna of New Mexico. 
Great Basin Naturalist 57: 283-314. 

Traver, J. R. 1938. Mayflies of Puerto Rico. Journal 
of Agriculture of the University of Puerto Rico 
22: 5-42. 

Waltz, R. D. and W. P McCafferty. 1987a. Generic 
revision of Cloeodes and description of two 
new genera (Ephemeroptera: Baetidae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 89: 177-184. 

. 1987b. Revision of the genus Cloeodes 

Traver (Ephemeroptera: Baetidae). Annals of 
the Entomological Society of America 80: 191- 
207. 

. 1994. Cloeodes (Ephemeroptera: Baetidae) 



in Africa. Aquatic Insects 16: 165-169. 



PROC. ENTOMOL. SOC. WASH. 

101(1). 1999. pp. 212-218 

NEW COMBINATIONS, NEW SYNONYMY, AND HYMONOMY IN THE 

ERIOCOCCIDAE, NEW HOMONOMY AND SYNONYMY IN THE 

CEROCOCCIDAE, AND TRANSFER OF CANCEROCOCCUS KOTEJA TO THE 

MARGARODIDAE (HEMIPTERA: COCCOIDEA) 

Douglass R. Miller and Maren E. Gimpel 

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, Bldg. 046, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: 
dmiller@sel.barc.usda.gov) 



Abstract. — A database and catalog of the eriococcid and cerococcid scale insects of the 
world is nearly complete and soon will be in press and placed on the World Wide Web. 
Before this is done, new combinations and other taxonomic changes need to be validated 
in print. This publication includes Neokaweckia Tang and Hao as a new synonym of 
Eriococcus, proposal of Neotrichococcus as a new name for Trichococcus Borchsenius, 
and new combinations in the family Eriococcidae; a new homonym and synonym in the 
Cerococcidae; and transfer of Cancerococcus from Eriococcidae to Margarodidae. 

Key Words: felt scales, Coccoidea, Eriococcidae, Cerococcidae, Margarodidae, ScaleNet, 
catalog, new combinations, Internet 



We currently are finishing a database and 
manuscript on the Eriococcidae and Cero- 
coccidae of the world. This research is part 
of a larger project called "ScaleNet" to de- 
velop a systematic database of the Coccoi- 
dea of the World; see Miller and Gimpel 
(1996), and Ben-Dov et al. (1997). One of 
the most controversial subjects in synthe- 
sizing the systematic data on eriococcids is 
to make sense out of the genera Acantho- 
coccus Signoret, Anophococcus Balachow- 
sky, Eriococcus Targioni Tozzetti, Gossy- 
paria Signoret, Greenisca Borchsenius, 
Gregoporia Danzig, Kaweckia Koteja and 
Zak-Ogaza, Neokaweckia Tang and Hao, 
and Rhizococcus Signoret. Most European 
literature recognizes all, or nearly all, of 
these genera as distinct, e.g., Kosztarab and 
Kozar (1988), but Hoy (1963) treated many 
of them as junior synonyms of Eriococcus 
and Williams (1985) treated all but Gre- 
goporia as members of Eriococcus. Gre- 



goporia was first treated as a synonym by 
Miller and Gimpel (1996) and Neokaweckia 
is here considered as a new junior synonym 
of Eriococcus. 

Although some have criticized the lump- 
ing of these genera as a reversion to Lin- 
naean times (Koteja 1997), we believe that 
the characters used to define these genera 
are homoplasious and discriminate artificial 
groups. It is logical to assume that natural 
groups occur in this assemblage, but it is 
important to undertake a careful phyloge- 
netic analysis to discover the groupings. 
Some cladistic work using molecular and 
morphological character systems is under- 
way in this regard by P. J. Gullan and Lyn 
Cook (Division of Botany and Zoology, 
Australian National University, Canberra). 
Their preliminary findings have been quite 
interesting and suggest the strong possibil- 
ity that there will be several genera within 
what is here treated as Eriococcus (Gullan, 



VOLUME 101, NUMBER 1 



213 



personal communication June 17, 1998). 
Unfortunately, this important research has 
been underway for only a short period of 
time and currently includes a small sample 
of the world eriococcid fauna. 

For many years, the first author surmised 
that the unusually large tubular ducts pre- 
sent on E. biixi (Fonscolombe) and E. eu- 
calypti Maskell were sufficient to charac- 
terize Eriococcus as separate from Acantho- 
coccus (see Miller and Williams 1976). 
However, it now appears that this is not 
necessarily the case and for the purposes of 
this paper we accept the conservative view 
pending results of the research by Gullan 
and her colleagues. This conservative view 
is consistent with the research of Ferris 
(1957), Hoy (1963), and WilUams (1985) 
and places all or most of the questionable 
genera in the genus Eriococcus. With this 
in mind, it is necessary to move several 
species previously placed in Acanthococcus 
into the genus Eriococcus for the first time. 

New Combinations in the Eriococcidae 

Eriococcus abaii (Danzig), n. comb. 

Acanthococcus abaii Danzig 1990: 
373. 
Eriococcus actius (Miller & Miller), n. 
comb. 

Acanthococcus actius Miller & Miller 
1993: 9. 
Eriococcus adzharicus (Hadzibejli), n. 
comb. 

Acanthococcus adzharicus Hadzibejli 
1960: 310. 
Eriococcus arenariae (Miller & Miller), n. 
comb. 

Acanthococcus arenariae Miller & 
Miller 1993: 13. 
Eriococcus barri (Miller), n. comb. 

Acanthococcus barri Miller 1991: 337. 
Eriococcus beshearae (Miller & Miller), n. 
comb. 

Acanthococcus beshearae Miller & 
Miller 1993: 15. 
Eriococcus brachypodii (Borchsenius & 
Danzig), n. comb. 



Greenisca brachypodii Borchsenius & 
Danzig 1966: 43. 
Eriococcus centaureae (Savescu), n. comb. 
Acanthococcus centaureae Savescu 
1985: 122. 
Eriococcus danzigae (Miller & Gimpel), n. 
comb. 

Rhizococcus confusus Danzig 1962a: 

854 (junior secondary homonym). 
Acanthococcus danzigae Miller & 
Gimpel 1996: 600 (replacement 
name). 

Remarks: The replacement name A. dan- 
zigae was given for Rhizococcus confusus 
Danzig (1962a) when it was transferred to 
Acanthococcus by Miller & Gimpel (1996) 
making it a junior secondary homonym of 
A. confusus (Maskell) (1892). The species 
epithet danzigae must continue to be used 
when transferred to Eriococcus, since A. 
confusus (Maskell) also is placed in Erio- 
coccus and is the senior homonym. 

Eriococcus davidsoni (Miller & Miller), n. 
comb. 

Acanthococcus davidsoni Miller & 
Miller 1993: 25. 
Eriococcus dennoi (Miller & Miller), n. 
comb. 

Acanthococcus dennoi Miller & Miller 
1993: 27. 
Eriococcus droserae (Miller, Liu, and How- 
ell), n. comb. 

Acanthococcus droserae Miller, Liu, 
and Howell 1992: 512. 
Eriococcus epacrotrichus (Miller & Mil- 
ler), n. comb. 

Acanthococcus epacrotrichus Miller & 
Miller 1992: 33. 
Eriococcus evelinae (Kozar), n. comb. 

Rhizococcus evelinae Kozar 1983: 

144. 
Acanthococcus evelinae (Kozar); Mill- 
er & Gimpel 1996: 600. 
Eriococcus froebeae (Miller), n. comb. 

Acanthococcus froebeae Miller 1991: 
343. 
Eriococcus herbaceus (Danzig), n. comb. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Rhizococciis herbaceus Danzig 1962b: 

22. 
Acanthococciis herbaceus; Nast et al. 

1990: 120. 
Acanthococcus herbaceus; Terezniko- 
va 1981: 29. 
Eriococcus hoyi (Miller & Miller), n. comb. 
Acanthococcus hoyi Miller & Miller 
1992: 44. 
Eriococcus iljiniae (Danzig), n. comb. 

Rhizococcus iljiniae Danzig 1972: 

339. 
Acanthococcus iljiniae; Miller & Gim- 
pel 1996: 601. 
Eriococcus istriensis (Kozar), n. comb. 

Gregoporia istriensis Kozar 1983: 

142. 
Acanthococcus istriensis; Miller & 
Gimpel 1996: 601. 
Eriococcus korotyaevi (Danzig), n. comb. 
Acanthococcus korotyaevi Danzig 
1982: 145. 
Eriococcus laeticoris (Tereznikova), n. 
comb. 

Greenisca laeticoris Tereznikova 

1965: 975. 
Kaweckia laeticoris; Koteja & Zak- 

Ogaza 1981: 507. 
Neokaweckia laeticoris; Tang & Hao 

1995: 514. 
Acanthococcus laeticoris; Miller & 
Gimpel 1996: 601. 
Eriococcus leptoporus (Miller & Miller), n. 
comb. 

Acanthococcus leptoporus Miller & 
Miller 1993: 39. 
Eriococcus niackenziei (Miller & Miller), n. 
comb. 

Acanthococcus niackenziei Miller & 
Miller 1992: 60. 
Eriococcus macrobactrus (Miller & Mil- 
ler), n. comb. 

Acanthococcus macrobactrus Miller & 
Miller 1992: 62. 
Eriococcus matesovae (Miller & Gimpel), 
n. comb. 

Acanthococcus multispinosus Mateso- 
va 1976: 24 (junior secondary hom- 
onym). 



Acanthococcus matesovae Miller & 
Gimpel 1996: 600 (replacement 
name). 

Remarks: The replacement name A. ma- 
tesovae was given for Acanthococcus mul- 
tispinosus Matesova (1976) when it was 
transferred to Acanthococcus by Miller & 
Gimpel (1996) making it a junior secondary 
homonym of A. multispinosus Kuhlgatz 
(1898). The species epithet matesovae must 
continue to be used when transferred to Er- 
iococcus, since A. multispinosus Kulgatz 
also is placed in Eriococcus and is the se- 
nior homonym. 

Eriococcus megaporus (Miller & Miller), n. 
comb. 

Acanthococcus megaporus Miller & 
Miller 1993: 45. 
Eriococcus mesotrichus (Miller & Miller), 
n. comb. 

Acanthococcus mesotrichus Miller & 
Miller 1993: 48. 
Eriococcus microtrichus (Miller & Miller), 
n. comb. 

Acanthococcus microtrichus Miller & 
Miller 1992: 65. 
Eriococcus minimus (Tang & Li), n. comb. 
Acanthococcus minimus Tang & Li 

1988: 71. 
Rhizococcus minimus; Tang & Hao 
1995: 352. 
Eriococcus monotrichus (Miller & Miller), 
n. comb. 

Acanthococcus microtrichus Miller & 
Miller 1993: 54. 
Eriococcus multispinatus (Tang & Hao), n. 
comb. 

Rhizococcus multispinatus Tang & 

Hao 1995: 598. 
Acanthococcus multispinatus; Miller 
& Gimpel 1996: 602. 
Eriococcus oligacanthus (Danzig), n. 
comb. 

Rhizococcus oligacanthus Danzig 

1972: 341. 
Acanthococcus oligacanthus; Miller & 
Gimpel 1996: 602. 



VOLUME 101, NUMBER 1 



215 



Eriococcus oligotrichus (Miller & Miller), 
n. comb. 

Acanthococcus oligotrichus Miller & 
Miller 1993: 57. 
Eriococcus ophius (Miller & Miller), n. 
comb. 

Acanthococcus ophius Miller & Miller 
1993: 59. 
Eriococcus orientalis (Borchsenius), n. 
comb. 

Greenisca orientalis Borchsenius 

1956: 676. 
Kaweckia orientalis (Borchsenius); 

Tang & Hao 1995: 511. 
Acanthococcus orientalis (Borchsen- 
ius); Miller & Gimpel 1996: 602. 
Eriococcus oxyacanthus (Danzig), n. comb. 
Acanthococcus oxyacanthus Danzig 

1975: 55. 
Rhizococcus oxyacanthus; Kozar & 
Walter 1985: 75. 
Eriococcus rubrus (Matesova), n. comb. 
Greenisca rubra Matesova 1960: 209. 
Kaweckia rubra; Koteja & Zak-Ogaza 

1981: 508. 
Neokaweckia rubra; Tang & Hao 

1995: 515. 
Acanthococcus rubra; Miller & Gim- 
pel 1996: 603. 
Eriococcus salicicola Tang, nomen nudum 

Remarks: Tang (1984) indicated that this 
species occurs widely over northeastern 
China on willow. He stated that he would 
be describing the species as new in the fu- 
ture, but there is no record of its publica- 
tion. 

Eriococcus stauroporus (Miller & Miller), 
n. comb. 

Acanthococcus stauroporus Miller & 
Miller 1992: 82. 
Eriococcus tosotrichus (Miller & Miller), n. 
comb. 

Acanthococcus tosotrichus Miller & 
Miller 1993: 62. 
Eriococcus washingtonensis (Miller & 
Miller), n. comb. 

Acanthococcus washingtonensis Miller 
& Miller 1992: 90. 



Eriococcus whiteheadi (Miller), n. 

comb. 
Acanthococcus whiteheadi Miller 

1991: 350. 
Eriococcus zernae (Tereznikova), n. comb. 
Acanthococcus zernae Tereznikova 

1977: 571. 

New Generic Synonymy in the 
Eriococcidae 

Neokaweckia Tang & Hao 1995: 596, new 
synonymy 

Type species: Greenisca rubra Matesova 
1960, by monotypy and original designa- 
tion 

Remarks: This genus is characterized by 
having a small anal ring, truncate body se- 
tae that are restricted to the last abdominal 
segments, and dorsal cruciform pores. 
These characters are considered to be with- 
in the expected range of variation for the 
genus Eriococcus. 

New Generic Homonomy and 
Replacement Name in the Eriococcidae 

Neotrichococcus Miller & Gimpel, new 
replacement name 

Trichococcus Borchsenius 1948: 503. 

Type species: Trichococcus filifer Borch- 
senius 1948, by monotypy and original des- 
ignation. 

Remarks: Trichococcus Borchsenius is a 
junior homonym of Trichococcus Kanda 
(1941) which is now considered to be a ju- 
nior synonym of Beesonia Green. Morrison 
and Morrison (1966) first discovered this 
homonymy but did not provide a replace- 
ment name. 

Neotrichococcus filifer (Borchsenius), n. 
comb. 

Trichococcus filifer Borchsenius 1948: 
503. 

Family Transfer of Cancerococcus to 

THE MaRGARODIDAE 

Cancerococcus Koteja 1988: 412, new 
family assignment 



216 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Remarks: This monotypic genus was 
originally placed in the Eriococcidae pre- 
sumably because of the enlarged setae on 
the antennae, a character found in many er- 
iococcid males. However, we have discov- 
ered that the genus is most closely related 
to the Pityococcini genera Pityococcus 
(McKenzie 1942) (including the species P. 
ferhsi McKenzie, P. deleoni McKenzie, P. 
rugulosus McKenzie) and Desmococcus (D. 
captivus McKenzie and D. sedentarius 
McKenzie) and is here transferred to the 
Margardodidae. The description of Cancer- 
ococcus (Koteja 1988) is based on a single 
wingless male from amber. The illustrations 
and description provided by Koteja were 
compared with a single undetermined male 
of Pityococcus deposited in the collection 
of National Museum of Natural History, 
Beltsville, Maryland. The following simi- 
larities occur in both taxa: Numerous short 
setae on the antennae; penial sheath that is 
apically bifurcate and has a broad aedeagus; 
antennae with short, round segments; eyes 
apparently numbering 4 (C. apterous) or 5 
{Pityococcus sp.) on each side of the head 
and set on a plate. This combination of 
characters is unique to the Pityococcini in 
the Margarodidae. 

New Synonymy in the Cerococcidae 

Asterococcus ramakrishnai (Ramakrishna 
Ayyar) 

Cerococcus ramakrishnae Ramachan- 
ran & Ramakrishna Ayyar 1934: 86. 
(nomen nudum) 

Cerococcus ramakrishnae Ramakrish- 
na Ayyar 1936: 148. 

Asterococcus ramakrishnai Lambdin 
1983: 304-306, new homonymy 
and synonymy. 

Remarks: The original combination of 
Cerococcus ramakrishnae is an unpub- 
lished manuscript name of Green. Rama- 
chanran & Ramakrishna Ayyar (1934) cited 
the name but gave no description thus cre- 
ating a nomen nudum. Ramakrishna Ayyar 
(1936) did not realize that he was validating 



the name and describing it after himself, but 
this was the case. Lambdin (1983) appar- 
ently knew of Green's manuscript name, 
but did not realize that Ramakrishna's de- 
scription was valid and described the spe- 
cies as new. Lambdin also corrected the 
spelling of the species epithet from "ra- 
makrishnae'' to ''ramakrishnai'' and 
moved the species from Cerococcus to As- 
terococcus. From the syntypes of Cerococ- 
cus ramakrishnae, we have chosen and 
marked as lectotype an adult female labeled 
in Green's handwriting: "on Ficus rootlets/ 
India, (Coimbatore)/ coll. Ramakrishna/ 24/ 
31. No. 335." (BMNH). The slide contains 
three specimens; the center specimen is the 
lectotype. There are three paralectotypes. 

Acknowledgments 

We thank the following individuals for 
their comments and suggestions relative to 
improvements of this manuscript: Douglas 
J. Williams, Department of Entomology, 
The Natural History Museum, London, UK 
and Michael E. Schauff, Systematic Ento- 
mology Laboratory, Agricultural Research 
Service, USDA, c/o National Museum of 
Natural History, Washington, DC. Funding 
for this paper was partially supported by 
BARD grant IS-2605. 95CR and is grate- 
fully acknowledged. 

Literature Cited 

Ben-Dov, Y. C. J. Hodgson, and D. R. Miller. 1997. 
Changes and comments on the taxonomy and no- 
menclature of some taxa in the families Coccidae, 
Eriococcidae and Pseudococcidae (Homoptera: 
Coccoidea). Phytoparasitica 25: 199-206. 

Borchsenius. N. S. 1948. On the revision of the genus 
Eriococcus Sign. (Insecta, Homoptera, Coccoi- 
dea). (In Russian). Doklady Akademii Nauk 
SSSR. Moscow (n.s.) 60: 501-503. 

. 1956. Notes on the Coccoidea of Korea. En- 

tomologicheskoe Obozrenye 35: 671-679. 

Borchsenius, N. S. and E. M. Danzig. 1966. A new 
species of Greenisca Borchs. (Homoptera, Coc- 
coidea, Eriococcidae) from the USSR, In Bykhov- 
skii, B. E., New Species of Insects of the Fauna 
of the USSR and Adjacent Countries. Trudy Aka- 
demii Nauk SSR Zoologicheskogo Instituta 37: 
41-44. 

Danzig, E. M. 1962a. Revision of the genus Rhizococ- 



VOLUME 101. NUMBER 1 



217 



cus Signoret (Homoptera, Coccoidea) of the SSR 
fauna. Entomologicheskoe Obozrenye 41: 839- 
860. 

. 1962b. Addition to the scale insect fauna (Ho- 
moptera, Coccoidea) of the Leningrad region. Tru- 
dy Akademii Nauk SSR Zoologicheskogo Insti- 
tuta. St. Petersburg 31: 22-24. 

. 1972. Contribution to the fauna of the white 

flies and scale insects (Homoptera: Aleyrodoidea, 
Coccoidea) of Mongolia. Insects of Mongolia 1: 
325-348. 

. 1975. Species of mealy-bugs (Homoptera, 

Coccoidea, Pseudococcidae) new for Mongolia. 
Insects of Mongolia 3: 48-55. 

. 1982. New species of the scale insects (Ho- 
moptera, Coccinea) from Mongolia. Insects of 
Mongolia 8: 140-147. 

. 1990. New species of coccids (Homoptera, 



Coccinea) from Iran, Mongolia and Vietnam. En- 
tomologicheskoe Obozrenye 69: 373-376. 

Ferris, G. F. 1957. A review of the family Eriococcidae 
(Insecta: Coccoidea). Microentomology 22: 81- 
89. 

Hadzibejli, Z. K. 1960. New species of coccids (Ho- 
moptera, Coccoidea) from Georgia. Trudy Aka- 
demii Nauk Gruzinskoy SSR Instituta Zashchitii 
Rastenii 13: 299-321. 

Hoy, J. M. 1963. A catalogue of the Eriococcidae (Ho- 
moptera: Coccoidea) of the world. New Zealand 
Department of Scientific and Industrial Research 
Bulletin 150: 1-260. 

Kanda, S. 1941. The genus Xylococcus and the genus 
Trichococcns [sic] N. G. from Japan (Homopteta) 
[sic]. Insect World 44: 68-72. 

Kosztarab, M. and F Kozar. 1988. Scale Insects of 
Central Europe. Akademiai Kiado, Budapest. 456 
pp. 

Koteja, J. 1988. Two new eriococcids from Baltic Am- 
ber Deutsche entomologische Zeitschrift 35: 405- 
416. 

. 1997. With Microsoft (through Windows) into 

the XVIII Century. Inclusion Wrostek 26: 17. 

Koteja, J. and B. Zak-Ogaza. 1981. Kaweckia gen. n. 
in the Eriococcidae (Homoptera, Coccoidea) and 
notes on related genera. Acta Zoologica Cracov- 
iensia 25: 501-518. 

Kozar, F. 1983. New and little-known scale-insect spe- 
cies from Yugoslavia (Homoptera: Coccoidea). 
Acta Zoologica Academiae Scientiarum Hungari- 
cae 29: 139-149. 

Kozar, F and J. Walter 1985. Check-list of the Pa- 
laearctic Coccoidea (Homoptera). Folia Entomo- 
logica Hungarica 46: 63-1 10. 

Kuhlgatz. 1898. Schildlause auf Kakteen nebst Be- 
schreibung von Rhizococciis multispinosus nov. 
spec. Monatsschrift fiir Kakteenkunde 8: 166- 
170; 185-188. 

Lambdin, P. L. 1983. A revision of the genus Astero- 



coccus Borchsenius (Homoptera: Cerococcidae). 
Proceedings of the Entomological Society of 
Washington 85: 297-308. 

Maskell, W. M. 1892. Further coccid notes: with de- 
scriptions of new species, and remarks on coccids 
from New Zealand, Australia and elsewhere. 
Transactions and Proceedings of the New Zealand 
Institute 24: 1-64. 

Matesova. G. I. 1960. New species of soft scales fam. 
Pseudococcidae (Homoptera, Coccoidea) of the 
Kazakhstan fauna. Trudy Instituta Zoologii, Aka- 
demii Nauk Kazakhskoy SSR, Alma-Ata 11: 205- 
217. 

1976. Two new species of eriococcids ("felt- 

coccoid") (Homoptera, Coccoidea, Eriococcidae) 
in Kazakhstan. Izvestiya Akademii Nauk Ka- 
zakhskoi SSR, (Seriya) Biologii 3: 22-26. 

McKenzie, H. L. 1942. New species of pine-infesting 
Margarodidae from California and southwestern 
United States (Homoptera; Coccoidea; Margarod- 
idae). (Contribution No. 30). Microentomology 7: 
1-18. 

Miller, D. R. 1991. Systematic analysis of Acantlio- 
coccus species (Homoptera: Coccoidea: Eriococ- 
cidae) infesting Atriple.x in western North Amer- 
ica. Proceedings of the Entomological Society of 
Washington 93: 333-355. 

Miller, D. R. and M. E. Gimpel. 1996. Nomenclatural 
changes in the Eriococcidae (Homoptera: Coccoi- 
dea). Proceedings of the Entomological Society of 
Washington 98: 597-606. 

Miller, D. R., T. Liu, and J. O. Howell. 1992. A new 
species of Acanthococcus (Homoptera; Coccoi- 
dea; Eriococcidae) from sundew (Drosera) with a 
key to the instars of Acanthococcus. Proceedings 
of the Entomological Society of Washington 
94(4): 512-523. 

Miller. D. R. and G. L. Miller. 1992. Systematic anal- 
ysis of Acanthococcus (Homoptera: Coccoidea: 
Eriococcidae) in the western United States. Trans- 
actions of the American Entomological Society 
118(1): 1-106. 

. 1993. Eriococcidae of the Eastern L^nited 

States (Homoptera). Contributions of the Ameri- 
can Entomological Institute 27(4): 1-91. 

Miller. D. R. and D. J. Williams. 1976. Proposed con- 
servation of the family-group name Eriococcidae 
Cockerell. 1899 (Insecta. Homoptera) and the des- 
ignation of a type-species for Eriococcus Targioni- 
Tozzetti. 1868 under the plenary powers Z.N.(S.) 
2140. Bulletin of Zoological Nomenclature 33: 
118-123. 

Morrison. H. and Morrison. E. R. 1966. An annotated 
list of generic names of the scale insects (Homop- 
tera: Coccoidea). Miscellaneous Publication Unit- 
ed States Department of Agriculture 1015: 1-206. 

Nast, J.. E. Chudzicka. S. M. Klimaszewski. W. Woj- 
ciechowski. Czylok. and J. Kotjea. 1990. [Check- 



218 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



list of Animals of Poland.] Wykaz zwierzat Polski. 
Razowski. J.. Ed. Zaklad Narodowy im. Ossolin- 
skich Wydawnictwo Polskiej Akademii Nauk 158 
pp. 

Ramachandran, S. and T V. Ramakiishna Ayyar 1934. 
Host plant index of Indo-Ceylonese Coccidae. 
Miscellaneous Bulletin, India Imperial Council of 
Agricultural Research 4: 1-113. 

Ramakrishna Ayyar, T. V. 1936. Notes on Coccidae 
(Homoptera, Rhynchota) from south India. Jour- 
nal of the Bombay Natural History Society 39: 
146-148. 

Savescu, A. D. 1985. Especes de coccoidees nouvelles 
pour la science signalees en Roumanie. III. Es- 
peces appartenant aux genres Pseiidococcus 
Westw., Phenacocciis Ckll., Paroudablis Ckll., 
Eupeliococciis Savescu et Lepidosaphes Shimer 
(Homoptera-Coccoidea). Bulletin de TAcademie 
des Sciences Agricoles et Forestieres. Bucarest 
14: 103-130. 

Tang, F T 1984. Observation on the scale insects in- 
jurious to forestry of North China. Shanxi Agri- 
cultural University Press Research Publication 2: 
122-133. 



Tang, F T and J. Hao. 1995. The Margarodidae and 
Others of China. Chinese Agricultural Science 
Technology Press Beijing, P. R. China, 738 pp. 

Tang. F T. and J. Li. 1988. Observations on the Coc- 
coidea of Inner Mongolia in China. Inner Mon- 
golia University Press, 227 pp. 

Tereznikova, E. M. 1965. New species of the genus 
Greenisca Borchs. (Coccoidea, Eriococcidae). 
Dopovidi Akademii Nauk Ukrainskoi RSR 7: 
957-959. 

Tereznikova, E. M. 1977. Two new species of Erio- 
coccidae from the genus of Acanthococcus Sign. 
(Homoptera, Coccoidea, Eriococcidae). Akademii 
Nauk Ukrains'koi RSR Dopovidi Seriya B 6: 
568-571. 

. 1981. Scale insects: Eriococcidae, Kermesi- 

dae and Coccidae. Fauna Ukraini. Akademiya 
Nauk Ukrainskoi RSR. Institut Zoologii. Kiev 20: 
1-215. 

Williams, D. J. 1985. The British and some other Eu- 
ropean Eriococcidae (Homoptera: Coccoidea). 
Bulletin of the British Museum (Natural History) 
Entomology Series 51: 347-393. 



PROC. ENTOMOL. SOC. WASH. 
101(1), 1999, pp. 219-220 



Note 



Scaphytopius angustatiis (Osbom) (Homoptera: Cicadellidae), a Leafhopper 
Characteristic of Pitch Pine-Scrub Oak Barrens 



Scaphytopius angustatiis (Osborn), a 
widely distributed Nearctic leafhopper of 
the deltocephaline tribe Scaphytopiini, is 
the only known conifer-feeding member of 
the genus (Hepner. 1947. University of 
Kansas Science Bulletin 31: 413-541). 
Adults are about 4 to 5 mm long, pale 
greenish fulvous or greenish yellow, with 
subhy aline fore wings; the distinctive male 
genitalia should be examined for positive 
identification (Hepner 1947; Berine. 1952. 
Canadian Entomologist 84: 311-313; 1956. 
Canadian Entomologist 88(Supplement 2): 
1-180). This pine specialist has been re- 
corded from jack pine {Pinus banks iana 
Lamb.), red pine (P. resinosa Alton), and 
pitch pine {P. rigida Mill.) (Ball. 1932. Ca- 
nadian Entomologist 64: 251-255; Hepner 
1947). Ecological information otherwise is 
lacking for this infrequently collected spe- 
cies. 

During studies of mirids (Wheeler. 1991. 
Journal of the New York Entomological So- 
ciety 99: 405-440) and fulgoroids (Wheeler 
and Wilson. 1996. Proceedings of the En- 
tomological Society of Washington 98: 
100-108) inhabiting pitch pine-scrub oak 
barrens, I found S. angustatiis to be a char- 
acteristic insect of northeastern pine bar- 
rens — that is, occurring consistently in, but 
not restricted to, that community type. It is 
one of several leafhopper species found on 
pitch pine in pine barrens. Individuals of 
the leafhopper were beaten from branches 
of pines, mainly pitch pine, as described by 
Wheeler (1991) for mirids occurring on 
scrub oak (Quercus ilicifolia Wangenh.). 
Voucher specimens have been deposited in 
the collections of Cornell University, Itha- 
ca, N.Y.; National Museum of Natural His- 
tory, Washington, D.C.; and the Pennsyl- 
vania Department of Agriculture, Harris- 
burg. 



Once considered a "distinctly northern 
species" (DeLong. 1923. pp. 56-163 In 
Britton, W.E., ed.. The Hemiptera or Suck- 
ing Insects of Connecticut. Connecticut 
Geological and Natural History Survey 
Bulletin 34), S. angustatiis is now known 
as far south as Georgia. Other records in- 
clude Maine, Massachusetts, Minnesota, 
Missouri, New Hampshire, New Jersey, 
New York, North Carolina, Ohio, Ontario, 
Pennsylvania, South Carolina, Virginia, and 
Wisconsin (Metcalf. 1967. pp. 2,075-2,695 
In General Catalogue of the Homoptera, 
Fascicle VI, Part 10, USDA ARS, Wash- 
ington, D.C.). Metcalf (1967) also listed 
Connecticut, but in a treatment of that 
state's leafhopper fauna, S. angustatus was 
only predicted to be found there (DeLong 
1923). 

During 1991-1995, I collected S. angus- 
tatus at 21 sites, ranging from extensive 
pitch pine-scrub oak barrens, such as Wa- 
terboro in Maine, Ossipee in New Hamp- 
shire, New York's Shawangunk Mountains, 
and the New Jersey Pine Barrens, to de- 
graded, remnant pine barrens. Seven collec- 
tions involved scattered pitch pines in com- 
munities other than pine barrens. Collec- 
tions were made from pitch pine except in 
the Gadway Barrens, Clinton Co., N.Y., 
where jack pine was the host. New state 
records are Connecticut, Rhode Island, and 
Vermont. Numbers in parentheses refer to 
adults unless otherwise stated. 

CONNECTICUT: Hartford Co., Shaker 
Pines, Enfield, 28 Sept. 1991 (3). MAINE: 
York Co., Biddeford, 14 Aug. 1993 (2); 
Kennebunk Plains, 13 Aug. 1993 (1); Wa- 
terboro Barrens Preserve, 8 Aug. 1995 (1, 
15 nymphs). MASSACHUSETTS: Franklin 
Co., Montague sand plains, 14 Sept. 1991 
(1), 15 Aug. 1993 (3, 1 nymph). NEW 



220 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



HAMPSHIRE: Carroll Co., Ossipee Pine 
Barrens, 7 Aug. 1995 (6 nymphs); Hills- 
borough Co., Amherst (1) & Nashua (1), 6 
Aug. 1995; Brookline, 14 Sept. 1991 (1). 
NEW JERSEY: Burlington Co., NW. of 
Warren Grove, 11 Aug. 1991 (2). NEW 
YORK: Albany Co., remnant barrens near 
Pine Bush Preserve, 22 Aug. 1993 (1); 
Clinton Co., NE. of Ausable Chasm, 30 
Aug. 1992 (1); Gadway Barrens, S. of Can- 
non Corners, 21 Aug. 1993 (1); West Chazy 
Banens, 29 Aug. 1992 (2); Jefferson Co., 
Plessis, 16 Aug. 1992 (>20 adults and 
nymphs); Saratoga Co., 3 mi. S. of South 
Glens Falls, 22 Aug. 1992 (1); Ulster Co., 
Mohonk Perserve near New Paltz, 29 Sept. 
1991 (2), 21 Aug. 1992 (3), 20 Aug. 1993 
(1). PENNSYLVANIA: Luzerne Co., Hum- 
boldt Industrial Park SW of Hazleton, 22 
Aug. 1993 (1); Schuylkill Co., near Frack- 
ville, 6 Oct. 1991 (1). RHODE ISLAND: 
Providence Co., Slatersville, 19 Sept. 1992 
(2); Washington Co., near Arcadia Manage- 
ment Area, 1 Sept. 1991 (2). VERMONT: 
Chittenden Co., Camp Johnson, Colchester, 
28 Aug. 1992 (1). 

Nymphs were seldom observed, and only 
the mostly bright green fifth instars were 
detected, earlier instars perhaps having been 
overlooked in the beating net. Because 
nymphs were not observed before August 
and no adults were taken during extensive 
spring and early-summer sampling of pitch 
pine, S. angustatus likely overwinters in the 
egg stage. 

Adults of this late-season, apparently 
univoltine leafhopper were observed in pine 
barrens from early August until early Oc- 
tober. In the northern part of its range (OH, 
NH, NY, WI), S. angustatus has been re- 
ported only from early August (Sanders and 
DeLong. 1917. Annals of the Entomologi- 



cal Society of America 10: 79-95) to mid- 
October (Osborn and Knull. 1947. Ohio 
Journal of Science 46: 329-336), even in 
New Hampshire, where Lowry (1933. Ohio 
Journal of Science 33: 59-80) collected 
leafhoppers nearly throughout the season, 
beginning in May. In the southern part of 
the range, adults have been collected in 
Georgia from June to August (Fattig. 1955. 
Emory University Museum Bulletin 11: 1- 
68). 

In northeastern pine barrens, I observed 
S. angustatus mainly on seedling and sap- 
ling pitch pines and the regrowth from 
stumps of cut-over trees. Certain other co- 
nifer-feeding leafhoppers are associated 
with seedlings, including those of pitch pine 
(DeLong, 1926. Ohio Journal of Science 
26: 69-72). About a third of the collections 
of S. angustatus were from mature pitch 
pines, where they often were beaten from 
bushy growth on the basal whorl of branch- 
es, some of which touched the ground. This 
growth habit, which might be a response to 
light reflected from the sand, typifies pitch 
pine in pine barrens (Kelley. 1927. Botan- 
ical Gazette 83: 89-93). 

I gratefully acknowledge those who ac- 
companied me in the field or enabled me to 
find pine barrens: Kenneth Adams, Robert 
Dirig, Paul Huth, Patrick McCarthy, Alan 
Nye, Dale Schweitzer, Nancy Sferra, and 
David VanLuven. Christopher Dietrich 
kindly identified S. angustatus. The Nature 
Conservancy and State Heritage Programs 
allowed access to several pine barrens, and 
Peter Adler provided useful comments on 
an early draft of the manuscript. 

A. G. Wheeler, Jr., Department of Ento- 
mology, Clemson University, Clemson, SC 
29634, U.S.A. (e-mail: awhlr@clemson.edu). 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 221-222 



Book Review 



Os Mosquitos de Macau (Diptera: Cu- 
licidae). Helena Cunha Ramos, Henrique 
Ribeiro, Maria Teresa Novo, and Emmett 
R. Easton. 1997. Sociedade Portuguesa 
de Entomologia, Apartado 8221, P-1800 
Lisbon. 201 pp., paper. ISBN 972-97241- 
0-5. 

Among the perquisites of my present po- 
sition is total access to the world literature 
on medical entomology. However, despite 
my databases, I am still occasionally sur- 
prised, as by the recent receipt of this lavish 
monograph on the mosquitoes of Macau 
(English Macao, Mandarin Aomen), Portu- 
gal's ancient entrepot on the South China 
Sea. At the eleventh hour, with Chinese of- 
ficials poised to take Government House, 
four widely experienced field biologists 
from the Centro de Zoologia, Instituto de 
Investiga^ao Cientifica Tropical (Cunha Ra- 
mos), the Disciplina de Entomologia Med- 
ica, Instituto de Higiene e Medicina Tropi- 
cal, Universidade Nova de Lisboa (Ribeiro 
and Teresa Novo), and the Centro de Es- 
tudos Pre-Universitarios, Universidade de 
Macau (Easton, an expatriate American 
profiled in J. New York Entomol. Soc. 102: 
389-391) have joined to forge a faunal sur- 
vey of continental proportions. 

Throughout 1994 and 1995, Cunha Ra- 
mos and colleagues scoured peninsular Ma- 
cau and its offshore islands of Taipa and 
Coloane, to which the cidade is linked by 
bridges and causeways. In an utterly an- 
thropocentric environment, they secured 
some 3,000 mosquito specimens represent- 
ing 28 species (generic and subgeneric ab- 
breviations follow Reinert, Mosq. Syst. 7: 
105-110, 14: 124-126, 23: 209-210): 

Aedes (Finlaya) togoi (Theobald 1907) 
Ae. (Stegomyia) albopictus (Skuse 1894) 
Ae. (Stg.) w-albus (Theobald 1905) 

Anopheles (Anopheles) sinensis Wiede- 
mann 1828 



Armigeres (Armigeres) subalbatus (Coquil- 
lett 1898) 
Ar. (Leicesteria) magnus (Theobald 
1908) 
Ciile.x (Ciile.x) bitaeniorhynchus Giles 1901 
Cx. (Cux.) jacksoni Edwards 1934 
Cx. (Cux. ) pseudovishnui CoWess 1957 
Cx. (Cux.) quinquefasciatus Say 1823 
Cx. (Cux.) sitiens Wiedemann 1828 
Cx. (Cux.) tritaeniorhynchus Giles 1901 
Cx. (Cux.) vagans Wiedemann 1828 
Cx. (Cux.) vishnui Theobald 1901 
Cx. (Culiciomyia) pallidothorax Theo- 
bald 1905 
Cx. (Eumelanomyia) foliatus Brug 1932 
Cx. (Eum.) ma lay i (Leicester 1908) 
Cx. (Lophoceraomyia) infantulus Ed- 
wards 1922 
Cx. (Lop.) ntbithoracis (Leicester 1908) 
Cx. (Lop.) sumatranus Brug 1931 
Cx. (Lutzia) fuscanus Wiedemann 1820 
Cx. (Lut.) halifaxii Theobald 1903 
Mansonia (Mansonioides) uniformis (Theo- 
bald 1901) 
Mimomyia (Mimomyia) chamberlaini Lud- 
low 1904 
To.xorhynchites (Toxorhynchites) macaensis 

Ribeiro 1997 
Tripteroides (Rachionotomyia) aranoides 

(Theobald 1901) 
Uranotaenia (Uranotaenia) annandalei 
Barraud 1926 
Ur. (Ura.) macfarlanei Edwards 1914 

Perusal of the regional literature (their bib- 
hography, pp. 195-201, contains 208 en- 
tries) yielded seven additional species: Ae. 
(Stg.) aegypti (Linnaeus 1762), An. (Cellia) 
jeyporiensis James 1902, A«. (Cel.) kanvari 
(James 1902), An. (Cel.) maculatus Theo- 
bald 1901, An. (Cel.) minimus Theobald 
1901, An. (Cel.) tessellatus Theobald 1901, 
and Cx. (Cux.) fuscocephala Theobald 
1907, all thought to occur in Macau in 
"very low densities." For a territorial total 
of 35 species, including the just-described 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



T. macaensis (J. Am. Mosq. Control Assoc. 
13: 213-217), keys are provided to males, 
females, and 4th instar larvae. The keys are 
exceptionally well supported with pen-and- 
ink drawings, paintings, and stunning color 
photomicrographs (of 179 numbered illus- 
trations, all but 46 are in full color). 

The core of the text (pp. 67-193) is a 
species-by-species account of mosquito tax- 
onomy, ecology, and distribution. Remark- 
able in this regard are three full-page, color- 
coded maps (pp. 68, 84 and 91) that pin- 
point collecting sites for each species on 
Taipa, Coloane, and Macau proper. Thus, it 
is possible to retrace the authors' steps to 
particular intersections, ponds, or cramped 
urban parks, where subsequent generations 
of the very insects that were the subjects of 
this study may yet be sought. As guidance 
to future investigators, photographs of 32 of 
these sites are included. The vector poten- 
tial of each species is also assessed for the 
malarias, 14 arboviruses, human filariases 
{Brugia malayi, Wuchereria bancrofti) and 
dirofilariases {Dirofilaria immitis, D. re- 
pens). 

A major goal of this endeavor was to 
compare the diversity of the Macanese 
mosquito fauna with that of neighboring 
Hong Kong and the Oriental Region as a 
whole. To this end, indices of abundance, 
association and distribution were calculated 
and are presented as tables throughout the 



text. These include the distribution of mos- 
quito species in Macau by principal juris- 
diction (Table 1); relative breeding indices 
for each species (Table 2); percentages for 
utilization of "natural" versus manmade 
habitats (Table 3); indices of larval associ- 
ation and their statistical significance (Ta- 
bles 4 and 5); the extraterritorial distribu- 
tion of each species by zoogeographic re- 
gion (Table 8); and comparative indices of 
biodiversity for the mosquito faunas of Ma- 
cau, Hong Kong, Cameroon, and mainland 
Portugal (Table 9). Such distillations be- 
speak countless hours of fieldwork — and 
countless more of analysis. 

The Portuguese are passing now. On 20 
December 1999, they will retrocede to Chi- 
na the oldest Western enclave in the Orient. 
How auspicious, for these two peoples have 
no quarrel with one another. The Lusitani- 
ans will be remembered not for foisting nar- 
cotics on a population held hostage, not for 
having to be forcibly evicted a la Dien Bien 
Phu, and certainly not for pushing the latest 
ludicrous cult from the so-called American 
heartland. Rather, both sides will be left to 
reflect on over 400 years of mutualism, as 
evidenced by this oblation. 

Richard G. Robbins, Armed Forces Pest 
Management Board, Walter Reed Army 
Medical Center, Washington, DC 20307- 
5001, U.S.A. 



PROC. ENTOMOL. SOC. WASH. 

101(1), 1999, pp. 222-224 



Book Review 



The Everglades Handbook: Under- 
standing the Ecosystem. Thomas E. 
Lodge, introduction by Marjory Stone- 
man Douglas. 1994, second printing, 
1998. St. Lucie Press, xix + 228 pp., pa- 
per. ISBN 1-884015-06-9. $39.95. 

The one person most closely associated 



with the formation of Everglades National 
Park was Marjory Stoneman Douglas. Her 
death earlier this year at 108 years of age 
marked the end of an important chapter in 
the history of south Florida. This book pro- 
vides an introduction to the Everglades, 
covering all aspects of the region. Mrs. 
Douglas' short introduction to the book 



VOLUME 101, NUMBER 1 



223 



serves as a brief memorial to her efforts to 
establish the Park. The Everglades Hand- 
book: Understanding the Ecosystem is writ- 
ten from an environmental protectionist 
perspective, which is to be expected of any 
work to which Mrs. Douglas contributed. 

The book is divided into four parts, each 
part being divided further into a number of 
chapters. The first part. Background, con- 
tains two chapters. Chapter 1, The Ever- 
glades in Space and Time, describes the 
geologic history of the Everglades. Chapter 
2, An Ecosystem Review, is an important 
chapter for the reader, for it is here that the 
author defines many terms referring to wa- 
ter and hydrology, including what the Ev- 
erglades are and are not, and more interest- 
ingly, where they are and where they are 
not. There is a great difference between 
what the Everglades historically were and 
what we now think of them as being. 

The second part of the book, Environ- 
ments of the Everglades Region, consists of 
seven chapters, each detailing one aspect of 
the Everglades. These chapters are weight- 
ed toward discussion of plant communities. 
Chapter 3 describes freshwater marshes, in- 
cluding the plants, soil, water, and effects 
of weather and fire on the marshes. Chapter 
4 discusses tree islands and their impor- 
tance to the region. Chapter 5 treats hard- 
wood hammocks and Chapter 6 pinelands. 
Both chapters discuss the role of fire in 
maintenance of the environment. Chapter 7 
is a very interesting discussion of mangrove 
swamps and their importance to wildlife 
and fisheries. Chapter 8 describes the veg- 
etation of the coastal lowlands and the ef- 
fects of hurricanes on south Florida. Final- 
ly, Chapter 9 deals with the estuaries and 
marine waters of the coast. This chapter dis- 
cusses the flora and geology of Florida Bay 
and the Gulf of Mexico, and the relation- 
ship between oysters and mangroves. 

Biogeography of Southern Florida, the 
third part of the book, probably will be of 
most interest to entomologists. There are 
eight chapters in this part of the book, seven 
of them covering a specific group of ani- 



mals. Chapter 10 includes such topics as the 
origin of the biota of southern Florida, and 
whether south Florida is tropical or sub- 
tropical. The Everglades region contains 
both temperate and tropical plant species, 
and the origins of each component of the 
flora are reviewed. Chapter 1 1 treats the in- 
vertebrates, marine, freshwater, and terres- 
trial. My bias as an entomologist led me to 
wonder why so few insect species were 
mentioned, but in fairness there is only one 
chapter to deal with all invertebrate ani- 
mals. Among the organisms mentioned are 
butterflies, crayfish, lobster, shrimp, snails, 
and spiders. Chapters 12 and 13 cover the 
fishes. Chapter 12 deals with freshwater 
species and Chapter 13 with estuarine and 
marine species. In Chapter 12 the distinc- 
tion among primary, secondary, and periph- 
eral freshwater fishes is explained. The im- 
portance of these fishes to the food chain is 
mentioned. Chapter 13 summarizes the di- 
versity of the marine and estuarine fishes, 
and their importance both as game fishes 
and as a food source for birds. Chapter 14 
is a very brief (three and one-half pages) 
mention of the amphibians of the Ever- 
glades. One interesting facet of this chapter 
is the impact that introduced frogs and 
toads have had on the native fauna. Chapter 
15 considers the reptiles. Most of the chap- 
ter deals with crocodilians, although lizards, 
snakes, tortoises, and turtles are mentioned. 
Chapter 16 is another very brief chapter 
that deals with mammals. Almost one entire 
page is devoted to the Florida panther. Oth- 
er species receive little attention. The last 
chapter in this part of the book. Chapter 17, 
discusses the birds. This chapter devotes 
much of its space to the large wading birds, 
which the author admits "attract much at- 
tention," and so they do here as well. This 
is one of the longer chapters in the book, 
comprising 18 pages. There is only brief 
mention of passerine birds, raptors, and oth- 
er types of birds. 

The fourth part of the book. Environ- 
mental Impacts, contains only one chapter. 
Chapter 18, that describes the effects that 



224 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



south Florida's increasing population has 
had on the Everglades. Specimen collect- 
ing, off-road vehicles, and introduced spe- 
cies are all mentioned as having adverse 
impacts on the Everglades. A great portion 
of the chapter is devoted to the effect that 
water control projects have had on the Ev- 
erglades. There is an interesting attempt to 
bring the global warming controversy into 
the discussion. However, the author does 
point out that there is another opinion with- 
in the scientific community, that of global 
cooling. The book ends with speculation on 
the demise of the Everglades. 

The book is extensively footnoted, and 
contains almost 300 references. The index 
is superb, permitting the reader to locate 
passages pertinent to any animal or plant by 
common or scientific name. The illustra- 
tions are of good quality. One disappoint- 
ment is that all photos are in black-and- 
white. The author writes in his preface that 
it was his intention to include a large num- 
ber of color photographs, but this became 
impossible due to realities of the publishing 
business. Many of the photos in the book 
were taken by photographer Robert Hamer, 
and their quality and composition leaves 



one hoping that the color version will some 
day see print. The shorter chapters are a bit 
frustrating to read, because the reader is 
given only enough information to whet the 
appetite. The shortness of some of the chap- 
ters apparently is due to their having been 
intended originally to accompany the color 
photos that never were used. The book was 
not conceived as a textbook, and it is evi- 
dent that the author did not intend it to be 
used as one. Readers who want a mathe- 
matical treatise detailing the population dy- 
namics of all species in the Everglades will 
not find that here. For those individuals 
who have an organismal interest in the Ev- 
erglades, e.g., naturalists and professional 
biologists in south Florida, the book will 
serve as a convenient introduction to some 
of the more spectacular animals and plants 
found in the region. For others, the book 
makes for pleasant reading, but its coverage 
of the invertebrate fauna is too limited to 
be of much use to the professional ento- 
mologist. 

Lawrence J. Hribar, Monroe County 
Mosquito Control District, Marathon, FL 
33050 U.S.A. 



PROC. ENTOMOL. SOC. WASH. 
101(1). 1999. p. 224 



1999 Meetings of the Entomological Society of Washington 



The following programs are scheduled 
for February to May, 1999. Meetings are at 
7:30 PM, the first Thursday of each month, 
in the National Museum of Natural History, 
Smithsonian Institution, Washington, DC. 

February 4, 1999 — Alan Schroeder (U.S. 
AID), "U.S. AID'S Pest Management 
Activities in Africa." 

March 4, 1999 — Dale E Schweitzer (The 
Nature Conservancy, Port Norris, NJ), 



"Lepidoptera and Other Insects in the 
New Jersey Pine Barrens." 

April 1, 1999 — Ted R. Schultz (Smithson- 
ian Institution), "The Natural History 
of Fungus-Growing Ants." 

May 6, 1999 — Joseph V. McHugh (Univer- 
sity of Georgia), "A Phylogenetic 
Analysis of Erotylidae (Coleoptera) 
with Implications for the Evolution of 
Their Mycophagy." 



PUBLICATIONS FOR SALE BY THE 
ENTOMOLOGICAL SOCIETY OF WASHINGTON 

Miscellaneous Publications 

A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. 

Schauff. 85 pp. 1990 $10.00 

A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric 

Grissell and Michael E. Schauff. 87 pp. 1997 15.00 

Memoirs of the Entomological Society of Washington 

Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. 

No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 $15.00 

No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 15.00 

No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 15.00 

No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 

Takahasi. 230 pp. 1969 15.00 

No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- 

pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 12.00 

No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 

67 pp. 1984 5.00 

No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 

pp. 1995 25.00 

No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent 

and W. L. Grogan, Jr. 198 pp. 1995 25.00 

No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 .. 25.00 

No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited 

by B. B. Norden and A. S. Menke. 216 pp. 1996 25.00 

No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and 

William L. Grogan, Jr. 297 pp. 1997 25.00 

No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by 

Thomas J. Henry. 149 pp. 1997 18.00 

No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- 

gafelter. 118 pp. 1998 12.00 

No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. 

Furth. 110 pp. 1998 12.00 

No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: 

Trichogrammatidae), by John D. Pinto. 287 pp. 1999 28.00 

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CONTENTS 

{Continued from front cover) 

KETH, A. C. and S. C. HARRIS — Two new species of Agarodes Banks (Trichoptera: Sericos- 

tomatidae) from southeastern United States 86 

KROMBEIN, KARL V. and ARKADY S. LELEJ — Biosystematic studies of Ceylonese wasps, XXII: 

Bethsmyrmilla, a new genus of mutillid wasps (Hymenoptera: Mutillidae: Myrmillinae) 143 

LUGO-ORTIZ, C. R., W. P. McCAFFERTY, and J.-L. GATTOLLIAT— The small minnow 

mayfly genus Cloeodes Traver (Ephemeroptera: Baetidae) in Madagascar 208 

MILLER, DOUGLASS R. and MAREN E. GIMPEL — New combinations, new synonymy, and 
hymonomy in the Eriococcidae, new homonomy and synonymy in the Cerococcidae, and 
transfer of Cancerococcus Koteja to the Margarodidae (Hemiptera: Coccoidea) 212 

SCHAEFER, CARL W. — ^The higher classification of the Alydidae (Hemiptera: Heteroptera) 94 

SITES, ROBERT W. and BECKY J. NICHOLS— Egg architecture of Naucoridae (Heteroptera): 

Internal and external structure of the chorion and micropyle I 

SLATER, JAMES A. and ALEX SLATER — Ashlockobius, a new genus of Myodochini from 

Venezuela (Hemiptera: Lygaeoidea: Rhyparochromidae: Myodochini) 138 

TROSTLE, M., N. S. CARREJO, I. MERCADO and R. A. WHARTON— Two new species of 

Phaenocarpa Foerster (Hymenoptera: Braconidae: Alysiinae) from South America 197 

NOTE 

WHEELER, A. G., JR. — Scaphytopius angustatus (Osborn) (Homoptera: Cicadellidae), a leaf- 
hopper, characteristic of pitch pine-scrub oak barrens 219 

BOOK REVIEWS 

ROBBINS, RICHARD G.—Os Mosquitos de Macau (Diptera: Culicidae) by H. C. Ramos, H. 

Ribeiro, M. T. Novo, and E. R. Easton 221 

HRIBAR, LAWRENCE J. — The Everglades Handbook: Understanding the Ecosystem, by Thomas 

E. Lodge 222 

MISCELLANEOUS 

1999 Meetings 224 



', 



» 



VOL. 101 



APRIL 1999 



NO. 2 

(ISSN 0013-8797) 




PROCEEDINGS 

of the 

ENTOMOLOGICAL SOCI 

of WASHINGTON 

PUBLISHED 
QUARTERLY 




CONTENTS 

ADAMSKI, DAVID — Two Neotropical Hypatopa Walsingham (Gelechioidea: Coleophoridae: 

Blastobasinae) with retractile labial palpi: A previously unknown lepidopteran feature 438 

BATRA, SUZANNE W. T. — Native bees (Hymenoptera: Apoidea) in native trees: Nyssa syl- 

vatica Marsh. (Cornaceae) 449 

CONTRERAS-RAMOS, ATILANO — List of species of Neotropical Megaloptera (Neuropterida) . . 274 

DIETRICH, CHRISTOPHER H.. MARK J. ROTHSCHILD, and LEWIS L. DEITZ— Checklist 

and host plants of the treehoppers (Hemiptera: Membracidae) of North Carolina 242 

FREIDBERG, AMNON — A new species of Craspedoxantha Bezzi from Tanzania and a revised 

phylogeny for the genus (Diptera: Tephritidae) 382 

GAGNE, RAYMOND J. and JOHN T. LILL — A new Nearctic species of Lestodiplosis (Diptera: 
Cecidomyiidae) preying on an oak leaf tier, Psilocorsis quercicella (Lepidoptera: Oeco- 
phoridae) 332 

GOEDEN, RICHARD D. and JEFFREY A. TEERINK— Life history and description of imma- 
ture stages of Trupanea wheeleri Curran (Diptera: Tephritidae) on Asteraceae in southern 
California 414 

HATHORNE, KEVIN T. and PATRICIA A. ZUNGOLI— Identification of late-instar nymphs 

of cockroaches (Blattodea) 316 

KIMSEY, LYNN S. — A turn of the century conundrum — reexamination of Aeolothynnus Ash- 
mead (Hymenoptera: Tiphiidae: Thynninae) 263 

KONDRATIEFF, BORIS C. and RICHARD W. BAUMANN— Studies on stoneflies of North 

Dakota with the description of a new Perlestci species (Plecoptera: Perlidae) 325 

KORCH, PETER P., STEVEN L. KEFFER, and ELISA WINTER— Description of immature 

stages of Platyvelia brachialis (Stal) (Heteroptera: Veliidae) 372 

LI, CHUN-LIN and PING-SHIH YANG— A new genus Clypeolontha Li and Yang, for the 
genus Melolontha Fabricius (Coleoptera: Scarabaeoidea: Melolonthinae) from southeastern 
Asia : 391 



(Continued on back cover) 



THE 

ENTOMOLOGICAL SOCIETY 

OF WASHINGTON 

Organized March 12, 1884 

OFFICERS FOR 1999 

Michael E. Schauff, President Michael G. Pogue, Treasurer 

David G. Furth, President-Elect John W. Brown, Program Chair 

Stuart H. McKamey, Recording Secretary Steven W. Lingafelter, Membership Chair 

HoLLis B. Williams, Corresponding Secretary Warren E. Steiner, Past President 
Andrew S. Jensen, Custodian 

David R. Smith, Editor 

Publications Committee 
Thomas J. Henry Wayne N. Mathis 

Gary L. Miller, Book Review Editor 



Honorary President 
Louise M. Russell 

Honorary Members 
Karl V. Krombein Ronald W. Hodges 



All correspondence concerning Society business should be mailed to the appropriate officer at the following 
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ington, D.C. 20560-0168. 

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tion, on the first Thursday of each month from October to June, inclusive, at 7:30 P.M. Minutes of meetings 
are published regularly in the Proceedings. 

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PLEASE SEE PP. 599-600 OF THE JULY 1998 ISSUE FOR INFORMATION REGARDING 
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STATEMENT OF OWNERSHIP 

Title of Publication: Proceedings of the Entomological Society of Washington. 

Frequency of Issue: Quarterly (January, April, July, October). 

Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of 
Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- 
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Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, 
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Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- 
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PRINTED BY ALLEN PRESS, INC.. LAWRENCE. KANSAS 66044. USA 

@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999. pp. 225-232 

DESCRIPTION OF IMMATURE STAGES OF TRUPANEA IMPERFECTA 
(COQUILLETT) (DIPTERA: TEPHRITIDAE) 

Jeffrey A. Teerink and Richard D. Goeden 

Department of Entomology, University of California, Riverside, CA 92521, U.S.A. 
(e-mail: rgoeden@ucracl.ucr.edu) 



Abstract. — The egg, first-, second- and third-instar larvae, and puparium of Tnipanea 
imperfecta (Coquillett), a monophagous, bi- or trivoltine tephritid principally reproducing 
in flower heads of Bebbia juncea (Bentham) Greene (Asteraceae) in southern California, 
are described and figured for the first time. The egg pedicel is composed mainly of a 
single row of large aeropyles. As with the other Tnipanea species previously studied, the 
lateral spiracular complex of the third instar is unique to T. impeifecta, with a stelex 
sensillum and two verruciform sensilla on the metathorax, and two verruciform sensilla 
on the abdominal segments. The third instar of T. imperfecta very closely resembles T. 
arizonensis Malloch in general habitus and sensory structures. 

Key Words: Insecta, Tnipanea, Asteraceae, nonfrugivorous Tephritidae, taxonomy of 
immature stages, egg, larva, puparium 



The life history of Tnipanea imperfecta 
(Coquillett) (Diptera: Tephritidae) was de- 
scribed by Goeden (1988) before adoption 
of our current format incorporating descrip- 
tion of the immature stages. To correct this 
deficiency and allow full comparison with 
the 36 species of southern California non- 
frugivorous fruit flies for which life histo- 
ries and descriptions of the immature stages 
have now been published, this paper de- 
scribes the immature stages of T. imperfec- 
ta. 

Materials and Methods 

One-liter samples of excised, immature 
and mature flower heads from the main host 
of T. imperfecta, Bebbia juncea (Bentham) 
Greene (Asteraceae), potentially containing 
eggs, larvae, and puparia were transported 
in cold-chests in an air-conditioned vehicle 
to the laboratory and stored under refrig- 
eration for subsequent dissection, photog- 
raphy, description, and measurement. 



Twenty-two eggs, 23 first-, 14 second-, and 
nine third-instar larvae, and nine puparia 
dissected from flower heads were preserved 
in 70% EtOH for scanning electron micros- 
copy (SEM). Specimens for SEM were hy- 
drated to distilled water in a decreasing se- 
ries of acidulated EtOH. They were osmi- 
cated for 24 h, dehydrated through an in- 
creasing series of acidulated EtOH and two, 
1-h immersions in Hexamethlydisilazane 
(HMDS), mounted on stubs, sputter-coated 
with a gold-palladium alloy, and studied 
with a Philips XL30-FEG SEM in the In- 
stitute of Geophysics and Planetary Phys- 
ics, University of California, Riverside. 

Plant names used in this paper follow 
Munz (1974); tephritid names follow Foote 
et al. (1993). Terminology and telegraphic 
format used to describe the immature stages 
follow Knio et al. (1996), Goeden and Teer- 
ink (1997, 1998, 1999), Goeden et al. 
(1998a, b), and Teerink and Goeden (1998), 
and our earlier works cited therein. Means 



226 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



± SE are used throughout this paper. 
Voucher specimens of T. imperfecta eggs, 
larvae and puparia are stored in a collection 
of immature Tephritidae acquired by JAT 
and now maintained by RDG. 

Results and Discussion 
Taxonomy 

Immature stages. — The egg and pupari- 
um of T. imperfecta were described and 
photographs of these stages and larvae were 
provided by Goeden (1988), but detailed 
descriptions based on scanning electron mi- 
croscopy heretofore have not been pub- 
lished. 

Egg: The egg (Fig. lA) of T. imperfecta 
has a short pedicel circumscribed by a sin- 
gle row of subrectangular aeropyles so 
large that they uniquely occupy more than 
half of this structure (Fig. IB). The micro- 
pyle is located on the anterior end of the 
pedicel (Fig. lC-1). 

First instar: White, elongate-cylindrical, 
rounded anteriorly and posteriorly, minute 
acanthae circumscribe intersegmental lines 
(Fig. 2A); gnathocephalon smooth, lacking 
rugose pads (Fig. 2C); dorsal sensory organ 
a dome-shaped papilla (Fig. 2B-1); subdor- 
sal sensillum located laterad of dorsal sen- 
sory organ (Fig. 2B-2); anterior sensory 
lobe bears terminal sensory organ (Fig. 2B- 
3), lateral sensory organ (Fig. 2B-4) and su- 
pralateral sensory organ (Fig. 2B-5); stomal 
sense organ ventrad of anterior sensory 
lobe (Fig. 2C-1); mouth hooks bidentate 
(Fig. 2C-2); median oral lobe laterally flat- 
tened (Fig. 2C-3); a pair of integumental 
petals dorsad of mouth hooks (Fig. 2C-4); 
pit sensillum laterad of mouth lumen (Fig. 
2C-5); minute acanthae ventrad of mouth 
lumen (Fig. 2C-6); anterior spiracle absent; 
abdominal lateral spiracular complex con- 
sists of a spiracle and two verruciform sen- 
silla; caudal segment with two stelex sen- 
silla dorsad and ventrad of posterior spirac- 
ular plates (Fig. 2D-1); two verruciform 
sensilla dorsolaterad of posterior spiracular 
plates (Fig. 2D-2); posterior spiracular plate 




Fig. 1. Egg of Tnipanea impetfecta: (A) habitus, 
pedicel to left; (B) egg pedicel; (C) pedicel, anterior 
view, 1 — micropyle, 2 — aeropyle. 



bears two ovoid rimae, ca. 0.008 mm in 
length (Fig. 2D-3), and four interspiracular 
processes, each with 1-3 branches, longest 
measuring 0.010 mm (Fig. 2D-4); inter- 
mediate sensory complex consists of a ste- 
lex sensillum (Fig. 2D-5) and a medusoid 
sensillum (Fig. 2D-6). 






VOLUME 101, NUMBER 2 



227 




Fig. 2. First instar of Tnipanea impeifecta: (A) habitus, anterior end to right; (B) anterior sensory lobe, 1 — 
dorsal sensory organ, 2 — subdorsal sensillum, 3 — terminal sensory organ, 4 — lateral sensory organ. 5 — supra- 
lateral sensory organ; (C) gnathocephalon, anterior view, 1 — stomal sense organ, 2 — mouth hook, 3 — median 
oral lobe, A — integumental petal, 5 — pit sensillum, 6 — minute acanthae; (D) caudal segment, 1 — stelex sensillum, 
2 — verruciform sensillum, 3 — rima, 4 — interspiracular process, 5 — intermediate sensory complex, stelex sensil- 
lum. 6 — intermediate sensory complex, medusoid sensillum. 



Second instar: White, elongate-cylindri- 
cal, tapering anteriorly, rounded posteriorly, 
minute acanthae circumscribe intersegmen- 
tal lines (Fig. 3A); gnathocephalon conical; 
rugose pads laterad of anterior sensory lobe 
(Fig. 3B-1); dorsal sensory organ a dome- 
shaped papilla (Fig. 3B-2, 3C-1); anterior 
sensory lobe bears terminal sensory organ 
(Fig. 3C-2), pit sensory organ (Fig. 3C-3), 
lateral sensory organ (Fig. 3C-4), and su- 
pralateral sensory organ (Fig. 3C-5); stomal 
sense organ ventrolaterad of anterior sen- 
sory lobe (Fig. 3B-3, 3C-6); mouth hooks 
bidentate (Fig. 3D-1); median oral lobe lat- 
erally flattened (Fig. 3D-2); labial lobe at- 
tached to median oral lobe (Fig. 3D-3); six 
pit sensilla circumscribe gnathocephalon 



(Fig. 3B-4); minute acanthae circumscribe 
anterior margin of prothorax (Fig. 3E-1); 
rugose pads (Fig. 3E-2) and two rows of 
verruciform sensilla circumscribe prothorax 
(Fig. 3E-3); anterior thoracic spiracles bear 
3 rounded papillae (Fig. 3E-4); lateral spi- 
racular complex not seen; caudal segment 
with minute acanthae dorsally (Fig. 3F-1); 
two stelex sensilla dorsad and ventrad of 
posterior spiracular plates (Fig. 3F-2); two 
verruciform sensilla dorsolaterad of poste- 
rior spiracular plates (Fig. 3F-3); posterior 
spiracular plate bears three ovoid rimae, ca. 
0.021 mm in length (Fig. 3F-4), and four 
interspiracular processes, each with 1-2 
branches, longest measuring 0.013 mm 
(Fig. 3F-5); intermediate sensory complex 



228 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



.fz^^^'^^^^^m^^^m- 




Fig. 3. Second instar of Trupanea imperfecta: (A) habitus, anterior to left; (B) gnathocephalon, anterior 
view. 1 — rugose pad, 2 — dorsal sensory organ, 3 — stomal sense organ, 4 — pit sensillum; (C) anterior sensory 
lobe, 1 — dorsal sensory organ, 2 — terminal sensory organ, 3 — pit sensory organ, 4 — lateral sensory organ, 5 — 
supralateral sensory organ, 6 — stomal sense organ; (D) gnathocephalon, ventral view, 1 — mouth hooks, 2 — 
median oral lobe, 3 — labial lobe; (E) gnathocephalon, prothorax, anterolateral view, 1 — minute acanthae, 2 — 
rugose pads, 3 — verruciform sensillum, 4 — anterior thoracic spiracle; (F) caudal segment, 1 — minute acanthae, 
2 — stelex sensillum, 3 — verrucifonn sensillum, 4 — rima. 5 — interspiracular process, 6 — intermediate sensory 
complex, medusoid sensillum. 7 — intermediate sensory complex, stelex sensillum. 



VOLUME 101. NUMBER 2 



229 



consisting of a medusoid sensillum (Fig. 
3F-6) and a stelex sensillum (Fig. 3F-7). 

Third instar: White, elongate-cylindri- 
cal, tapering anteriorly, rounded posteriorly, 
minute acanthae circumscribe intersegmen- 
tal lines (Fig. 4A); gnathocephalon conical 
(Fig. 4B), rugose pads laterad of anterior 
sensory lobe (Fig. 4B-1), those laterad of 
mouth lumen serrated on ventral margin 
(Fig. 4C-1); dorsal sensory organ a dome- 
shaped papilla (Fig. 4B-2, 4C-2); subdorsal 
sensillum laterad of dorsal sensory organ; 
anterior sensory lobe (Fig. 4B-3, 4C) bears 
terminal sensory organ (Fig. 4C-3), pit sen- 
sory organ (Fig. 4C-4), lateral sensory or- 
gan (Fig. 4C-5), and supralateral sensory 
organ (Fig. 4C-6); stomal sense organ ven- 
trolaterad of anterior sensory lobe (Fig. 4B- 
4, 4C-7); mouth hooks tridentate (Fig. 4B- 
5); median oral lobe laterally flattened, ta- 
pering anteriorly (Fig. 4B-6); prothorax cir- 
cumscribed anteriorly with minute acanthae 
(Fig. 4D-1); rugose pads circumscribe pro- 
thorax posteriorad to minute acanthae (Fig. 
4D-2); two rows of verruciform sensilla cir- 
cumscribe prothorax posteriorad to rugose 
pads (Fig. 4D-3); stelex sensillum located 
dorsomedially (Fig. 4D-4); anterior thoracic 
spiracle bears three rounded papillae (Fig. 
4D-5); mesothorax and metathorax circum- 
scribed anteriorly with verruciform sensilla; 
metathoracic lateral spiracular complex 
consists of a spiracle (Fig. 4E-1), a stelex 
sensillum (Fig. 4E-2), and two verruciform 
sensilla (Fig. 4E-3); abdominal lateral spi- 
racular complex consists of a spiracle (Fig. 
4F-1) and two verruciform sensilla (Fig. 
4F-2); caudal segment circumscribed by 
minute acanthae; two stelex sensilla dorsad 
and ventrad of posterior spiracular plates 
(Fig. 4G-1); two verruciform sensilla dor- 
solaterad of posterior spiracular plates (Fig. 
4G-2); posterior spiracular plate bears three 
ovoid rimae, ca. 0.03 mm in length (Fig. 
4G-3), and four interspiracular processes, 
each with 2-4 branches, longest measuring 
0.02 mm (Fig. 4G-4); intermediate sensory 
complex consists of a medusoid sensillum 



(Fig. 4H-1), and a stelex sensillum (Fig. 
4H-2). 

Puparium: Black, elongate-cylindrical 
(Fig. 5A); anterior end bears the invagina- 
tion scar (Fig. 5B-1), and anterior spiracles 
(Fig. 5B-2); caudal segment circumscribed 
by minute acanthae (Fig. 5C-1), two stelex 
sensilla dorsad and ventrad of posterior spi- 
racular plates (Fig. 5C-2); two verruciform 
sensilla dorsolaterad of posterior spiracular 
plates (Fig. 5C-3); posterior spiracular plate 
bears three ovoid rimae (Fig. 5C-4), and 
four interspiracular processes, each with 2- 
4 branches (Fig. 5C-5); intermediate sen- 
sory complex consists of a medusoid sen- 
sillum and a stelex sensillum (Fig. 5C-6). 

Discussion 

The egg of Trupanea imperfecta is elon- 
gate-ellipsoidal, with a reduced peg-like an- 
terior pedicel (Goeden 1988). It is similar 
in size and shape to T. signata Foote (Goe- 
den and Teerink 1997), longer than T. ac- 
tinobola (Loew) (Goeden et al. 1998b), T. 
californica Malloch (Headrick and Goeden 
1991), and T. pseudovicina Hering (Goeden 
and Teerink 1998), and shorter but wider 
than T. arizonensis (Goeden and Teerink 
1999). The pedicel is mainly composed of 
large aeropyles (Figure IB, IC). The pedi- 
cels of 71 arizonensis, T. jonesi Curran, T. 
nigricornis (Coquillett), and T. pseudovici- 
na are similar in shape, but have smaller 
aeropyles spaced farther apart (Goeden and 
Teerink 1998, 1999; Goeden et al. 1998a; 
Knio et al. 1996). 

The first instar of T. imperfecta is similar 
in general habitus to previously studied 
Trupanea species (Goeden and Teerink 
1998, 1999; Goeden et al. 1998a, b; Knio 
et al. 1996; Teerink and Goeden 1998). The 
gnathocephalon is smooth, lacking rugose 
pads. The pit sensory organ is indistinct on 
the anterior sensory lobe, and the stomal 
sense organ is also indistinct. Minute acan- 
thae are limited to the ventral margin of the 
prothorax. The interspiracular processes, 
each with 1-3 blade-like processes, are sim- 
ilar to T. actinobola, T. bisetosa (Coquil- 



230 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 







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Fig. 4. Third instar of Trupauea impeifecta: (A) habitus, anterior to left; (B) gnathocephalon. anterior view, 
1 — rugose pads, 2 — dorsal sensory organ, 3 — anterior sensory lobe, 4 — stomal sense organ, 5 — mouth hook, 
6 — median oral lobe; (C) anterior sensory lobe, 1 — serrated rugose pad. 2 — dorsal sensory organ, 3 — terminal 
sensory organ, 4 — pit sensory organ, 5 — lateral sensory organ, 6 — supralateral sensory organ, 7 — stomal sense 
organ; (D) gnathocephalon, prothorax, anterior view, 1 — minute acanthae, 2 — rugose pad, 3 — verruciform sen- 
sillum, 4 — stelex sensillum, 5 — anterior thoracic spiracle; (E) metathorax, 1 — spiracle, 2 — stelex 



VOLUME 101, NUMBER 2 



231 




Fig. 5. Pupariuni of Triiponea impeifecta: (A) 
habitus, anterior end to left; (B) anterior end, 1 — in- 
vagination scar, 2 — anterior thoracic spiracle; (C) cau- 
dal segment, 1 — minute acanthae, 2 — stelex sensillum, 
3 — verruciform sensillum. 4 — rima, 5 — interspiracular 
process, 6 — intermediate sensory complex. 



lett), and T. pseudovicina (Goeden and 
Teerink 1998; Goeden et al. 1998b; Knio et 
al. 1996). 

The second instar of T. imperfecta differs 
from the first instar in possessing rugose 
pads laterad of the mouth lumen, and the 
anterior margin of the prothorax is circum- 
scribed by minute acanthae, rugose pads, 
and two rows of verruciform sensilla. All 
four of the anterior sensory lobe sensilla, as 
well as the stomal sense organ, are distinct 
in the second instar. The anterior thoracic 
spiracle is present in the second instar, and 
the posterior spiracular plates possess three 
ovoid rimae rather than two as in the first 
instar. The caudal segment in the second in- 
star, unlike in the first instar, is circum- 
scribed by minute acanthae. 

The third instar differs from the second 
instar in possessing serrated rugose pads 
laterad of the mouth lumen, and the mouth 
hooks tridentate. The third instar of T. im- 
perfecta is very similar in general habitus 
to T. arizonensis and T. pseudovicina, being 
more elongate-cylindrical than barrel- 
shaped (Goeden and Teerink 1998, 1999). 
Trupanea imperfecta and T. arizonensis are 
also similar, in that the meso- and metatho- 
rax are circumscribed by verruciform sen- 
silla and the anterior thoracic spiracle bears 
three ovoid papillae (Goeden and Teerink 
1998). However, there are slight differences 
between these two species in that the rugose 
pads laterad of the mouth lumen are senat- 
ed in T. imperfecta, but not in T. arizonen- 
sis. Moreover, the metathoracic lateral spi- 
racular complex is slightly different, with a 
stelex sensillum and two verruciform sen- 
silla in T. imperfecta, and three verruciform 
sensilla in T. arizonensis (Goeden and Teer- 
ink 1999). Trupanea pseudovicina is simi- 
lar to T. imperfecta in having serrated ru- 
gose pads laterad of the mouth lumen (Goe- 



sensillum, 3 — verruciform sensilla; (F) sixth abdominal segment, 1 — spiracle, 2 — verruciform sensilla: (G) cau- 
dal segment, 1 — stelex sensillum, 2 — verruciform sensillum, 3 — rima, 4 — interspiracular process; (H) interme- 
diate sensory complex, 1 — medusoid sensillum, 2 — stelex sensillum. 



232 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



den and Teerink 1998). It differs from T. 
impetfecta by having four, not three, papil- 
lae on the anterior spiracle; the meso- and 
metathorax not circumscribed by verruci- 
form sensilla, and a stelex sensillum and 
two, not one, verruciform sensilla in the 
metathoracic lateral spiracular complex 
(Goeden and Teerink 1998). As with the 
other Trupanea species studied by us to 
date, the lateral spiracular complex is 
unique to T. imperfecta (Goeden and Teer- 
ink 1997, 1998, 1999; Goeden et al. 1998a, 
b; Headrick and Goeden 1991; Knio et al. 
1996, Teerink and Goeden 1997). 

The puparium of T. imperfecta is larger 
than T. actinobola and T. californica (Goe- 
den et al. 1998b; Headrick and Goeden 
1991 ), wider but shorter than T. arizonensis 
and T. pseudovicina (Goeden and Teerink 
1998, 1999). 

Acknowledgments 

We are grateful to David Headrick for his 
helpful comments on an earlier draft of this 
paper. 

Literature Cited 

Foote. R. H., E L. Blanc, and A. L. Nonboni. 1993. 
Handbook of the Fruit Flies (Diptera: Tephritidae) 
of America North of Mexico. Cornell University 
Press. Ithaca, New York. 

Goeden, R. D. 1987. Life history of Trupanea con- 
jiincUi (Adams) on Trixis californica Kellogg in 
southern California (Diptera: Tephritidae). Pan- 
Pacific Entomologist 63: 284-291. 

. 1988. Life history of Trupanea inipeifecta 

(Coquillett) on Behhia juncea (Bentham) Greene 
in the Colorado Desert of southern California 
(Diptera: Tephritidae). Pan-Pacific Entomologist 
64: 345-351. 



Goeden, R. D. and J. A. Teerink. 1997. Life history 
and description of immature stages of Trupanea 
sif^nata Foote (Diptera: Tephritidae) on Gnaphal- 
ium hiteo-album L. in southern California. Pro- 
ceedings of the Entomological Society of Wash- 
ington 99: 748-755. 

. 1998. Life history and description of imma- 
ture stages of Trupanea pseudovicina Hering 
(Diptera: Tephritidae) on Porophyllum gracile 
Bentham (Asteraceae) in southern California. Pro- 
ceedings of the Entomological Society of Wash- 
ington 100: 361-372. 

. 1999. Life history and description of imma- 
ture stages of Trupanea arizonensis Hering (Dip- 
tera: Tephritidae) on Trixis californica Kellogg 
var. californica (Asteraceae) in southern Califor- 
nia. Proceedings of the Entomological Society of 
Washington 101: 75-85. 

Goeden, R. D., J. A. Teerink, and D. H. Headrick. 
1998a. Life history and description of immature 
stages of Trupanea jonesi Curran (Diptera: Te- 
phritidae) on native Asteraceae in southern Cali- 
fornia. Proceedings of the Entomological Society 
of Washington 100: 126-140. 

. 1998b. Life history and description of im- 
mature stages of Trupanea actinobola (Loew) 
(Diptera: Tephritidae) on Acamptopappus spliaer- 
ocephalus (Harvey and Gray) Gray (Asteraceae) 
in southern California. Proceedings of the Ento- 
mological Society of Washington 100: 674-688. 

Headrick. D. H. and R. D. Goeden. 1991. Life history 
of Trupanea californica Malloch (Diptera: Te- 
phritidae) on Gnaphalium spp. in southern Cali- 
fornia. Proceedings of the Entomological Society 
of Washington 93: 559-570. 

Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996. 
Descriptions of immature stages of Trupanea ni- 
gricornis and T. bisetosa (Diptera: Tephritidae) 
from southern California. Annals of the Entomo- 
logical Society of America 89: 1-11. 

Munz, P. A. 1974. A Flora of Southern California. Uni- 
versity of California Press, Berkeley and Los An- 
geles. 

Teerink, J. A. and R. D. Goeden. 1998. Description of 
immature stages of Trupanea conjuncta (Adams) 
(Diptera: Tephritidae). Proceedings of the Ento- 
mological Society of Washington 100: 431-438. 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999, pp. 233-241 

THE MICROCADDISFLY GENUS ITHYTRICHIA EATON (TRICHOPTERA: 
HYDROPTILIDAE) IN NORTH AMERICA 

Stephen R. Moulton II, Steven C. Harris, and Joseph P. Slusark 

(SRM, JPS) U.S. Geological Survey, National Water Quality Laboratory, Biological Unit, 
PO. Box 25046, MS 407, Denver, CO 80225, U.S.A. (e-mail, SRM: smoulto@usgs.gov); 
(SCH) Department of Biology, Claiion University, Clarion, PA 16214, U.S.A. 



Abstract. — The distribution and taxonomy of the microcaddisfly genus Ithytrichia Eaton 
in North America is reviewed. Males and females of /. clavata Morton, /. mazon Ross, 
and /. mexicana Harris and Contreras-Ramos are illustrated, and a key is provided for 
their separation. Females of /. mazon and /. mexicana are described for the first time; the 
female of /. clavata is redescribed. 

Key Words: Trichoptera, Hydroptilidae, Ithytrichia, taxonomy, nearctic distribution 



The holarctic genus Ithytrichia (Eaton 
1873) is a small group of microcaddisflies 
with six species worldwide (Marshall 1979, 
Morse 1993) belonging to the subfamily 
Hydroptilinae, tribe Orthotrichiini. Three 
species, /. clavata Morton, /. mazon Ross, 
and /. mexicana Harris and Contreras-Ra- 
mos, are found in North America (Morse 
1993). Adults are distinguished from those 
of other nearctic hydroptilid genera by the 
presence of ocelli, a 0-3-4 tibial spur count, 
mesoscutellum without transverse suture, 
and posterodorsal margin of mesoscutellum 
separated from posterior margin by a nar- 
row strap (Moulton and Stewart 1996). The 
laterally compressed abdomen having dor- 
sal and ventral membranous lobes easily 
identifies larvae of Ithytrichia; the larval 
case is composed entirely of silk and is 
purse-like with a small circular anterior 
opening (Wiggins 1996). Morphological 
characters have not been discovered to dis- 
tinguish the larvae to species. Before this 
study, only the female of /. clavata was 
known. In this paper, we describe for the 
first time the females of /. mazon and /. 
mexicana. Males of the three species are 



reillustrated with accompanying distribu- 
tional notes. Keys for separating males and 
females of the three North American spe- 
cies are also provided. 

Material examined in this study is depos- 
ited at the Arkansas State University Muse- 
um of Zoology, Jonesboro (ASUMZ), the 
California Academy of Sciences, San Fran- 
cisco (CAS), the C. P. Gillette Museum of 
Arthropod Diversity, Colorado State Univer- 
sity, Fort Collins (CSU), the Illinois Natural 
History Survey, Champaign (INHS), the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 
(NMNH), the Ohio Biological Survey, Co- 
lumbus (OBS), the University of Minnesota, 
St. Paul (UM), the University of North Tex- 
as, Denton (UNT), and in the research col- 
lection of the senior author (SRM). Al- 
though most specific characters may be dis- 
cerned by using a dissecting microscope 
(60-lOOX), it is necessary to view some of 
the female genitalic characters (e.g., sper- 
mathecal sclerite) by using a compound mi- 
croscope (100-400X). Morphological ter- 
minology follows that of Marshall (1979). 
Length is measured from the tip of the head 
to the posterior tip of the forewings. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Ithytrichia clavata Morton 
(Figs. 1, 4, 7) 

Ithytrichia clavata Morton 1905:67. 

Ross (1944) described the female of /. 
clavata, however, the discovery of the fe- 
males of /. mazon and /. mexicana neces- 
sitates a redescription for comparison. 

Female description. — Length 2.9-3.5 
mm. 21 antennal segments. Light brown in 
alcohol. Sternite VI with short acute ventro- 
mesal process. Sternite VII in ventral view 
dome-shaped. Sternite VIII in ventral view 
parallel-sided, posterior margin with series 
of stout setae, each arising from a membra- 
nous tubercle; ventral sclerite widening 
posteriorly, posterior margin with broad 
concavity, bi-lobed, each lobe with a mem- 
branous pocket; two pairs of lateral apo- 
demes. Segment IX bullet-shaped in ventral 
view; one pair of lateral apodemes. Seg- 
ment X button-like with a pair of short cer- 
ci. Apodemes slender and rod-like, one pair 
extending from anterior end of segment X 
apodemes, other pair from anterior edge of 
segment VIII, both pairs extending to seg- 
ment VI, with anterior apices gently curv- 
ing mesad. Spermathecal sclerite in ventral 
view with anterior membranous and circu- 
lar; basal one-third sclerotized on lateral 
margins, anterior apices angled laterad, 
gradually tapering posteriorly to pointed, 
incurved apices, middle portion of sclero- 
tized apparatus arrowhead-shaped; posterior 
two-thirds widest at base, narrowing in 
middle to tubular apex, middle portion with 
lateral patches of minute spines. 

Material examined. — USA: CALIFOR- 
NIA, Colusa Co., Bear Creek, 26 km E 
Clearlake Highlands, 28-VII-1974, P. Peter- 
son, 6 6 (CAS); Lake Co., elev. 402 m, 16- 
IX-1949, H. R Chandler, 1 S (CAS); Napa 
Co., Capell Cr., 7-VM952, 1 6 (CAS); Big 
Canyon Creek, 13 km NE Middletown, 23- 
VI- 1974, R Peterson, 1 6 (CAS); FLORI- 
DA, Jackson Co., Florida Caverns State 
Park, 4-V-1970, 2 6 (NMNH); ILLINOIS, 
Galena River, Council Hill, 26-VI-1940, 
Mohr and Riegel, 10 6 (INHS); MAINE, 



Ashland, 29-VII-1924, 1 S (CAS); Orono, 
Lake Pushaw, 1-3-VIII-1966, W. W. Wirth, 
3 6 (NMNH); Oxbow (T9R5), 22-Vn- 
1961, A. Brower, 2 6 (NMNH); Allagash, 
29-VII-1959, 32 6 (NMNH); same but, 30- 
VII- 1959, 130 6 (NMNH); same but, 5- 
VIII-1959, 8 6 (NMNH); MISSOURI, 
Gasconade Co., Gasconade River, Held's 
Island Access, 28-VIII-1990, B. C. Poulton, 
3 6 (UNT); Maries Co., Gasconade River, 
Island Ford Resort @ Hwy 42, 7-VIII, 
1990, B. C. Poulton, 1 S (UNT); same but, 
Paydown Access, SW Belle, 15-VII-1990, 
1 6 (UNT); Osage Co., Gasconade River, 
Hwy 89 @ Dallas Ferry Access, 27-IX- 
1990, B. C. Poulton, 2 6 (UNT); PENN- 
SYLVANIA, Presque Isle, 15-VIII-1947, 1 
6 (CAS); Chemung River, Athens, 8-VII- 
1937, J. Eddleston, 23 6 (INHS); TEXAS, 
Brewster Co., county park, 8 km S Mara- 
thon, 22-VI-1994, B. Kondratieff, 5 S, 
(CSU); Edwards Co., South Llano River @ 
Paint Rock Springs, Hwy 337, 12-VI-1992, 
B. Kondratieff, 1 S (CSU); Hays Co., Blan- 
co River @ Post Rd., 14-V-1991, S. Tie- 
mann, 3 6 (UNT); Palo Pinto Co., Brazos 
River, TX Hwy 4, 23-III-1972, Stark and 
Rhame, 1 (5, 1 9 (NMNH), same but, 6- 
VI-1995, D. C. Houghton, 27 S, 59 9 
(UNT); Randall Co., Prairie Dog Town 
Fork of Red River, Palo Duro Canyon State 
Park, water crossing No. 5, 12-IX-1997, S. 
R. Moulton and G. W. Easley, 2 6,19 
(SRM); WISCONISIN, Door Co., Egg Har- 
bor, 13-VIII-1940, C. O. Mohr, 21 6, 15 9 
(INHS); same but, Ephraim, WI, ?-?-1957, 
1 9 (INHS); CANADA: MANITOBA, 
Lake Manitoba, 5 km W Delta, 3-VIII- 
1967, D. Webb, 9 6 (INHS); QUEBEC, Ot- 
tawa River, Quyon, 1 -VIII- 1976, O. S. 
Flint, Jr., 4 d, 1 9 (NMNH); Norway Bay, 
4-VIII-1973, O. S. FHnt, Jr., 4 d, 4 9 
(NMNH). 

Discussion. — Ithytrichia clavata has a 
holarctic distribution (Fischer 1961); it is 
widely distributed throughout the United 
States and southern Canada (Wiggins and 
Parker 1997). Houghton and Stewart (1998) 
reported on the seasonal flight periodicity 



VOLUME 101. NUMBER 2 



235 




D 





Fig. 1. Ithythchia clavata. male genitalia. A, Left lateral. B, Dorsal. C. Ventral. D, Phallus. 



236 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



of /. clavata from the Brazos River in 
north-central Texas. Specimens of/, clavata 
from the eastern United States determined 
before 1944 should be checked to ensure 
they are not misidentifications of /. mazon. 
We found this to be true in some of the 
material we examined. 

Ithytrichia mazon Ross 
(Figs. 2, 5, 8) 

Ithytrichia mazon Ross 1944:124. 

Female description. — Length 3.3-4.0 
mm. 21 antennal segments. Light brown in 
alcohol. Sternite VI with short acute ventro- 
mesal process. Sternite VII in ventral view 
dome-shaped. Sternite VIII in ventral view 
parallel-sided, posterior margin with series 
of stout setae, each arising from a membra- 
nous tubercle; ventral sclerite widening 
posteriorly, apex truncate. Segment IX bul- 
let-shaped in ventral view, apex membra- 
nous. Segment X button-like with a pair of 
short cerci. Apodemes slender and rod-like, 
one pair extending from anterior end of seg- 
ment X apodemes, other pair from anterior 
edge of segment VIII, both pairs extending 
to segment VI, with anterior apices gently 
curving laterad. Spermathecal sclerite in 
ventral view with anterior a membranous 
funnel-shape; basal one-third sclerotized 
laterally and bowed, middle portion of 
sclerotized apparatus arrowhead-shaped; 
posterior two-thirds widest at base, narrow- 
ing in middle to tubular apex, middle por- 
tion with lateral patches of minute spines. 

Material examined. — USA: ARKAN- 
SAS, Logan Co., Sixmile Creek, 23-V- 
1986, H. W. Robison, 1 6 (INHS); ILLI- 
NOIS, Mazon, along Mazon Creek, 16-VI- 
1938, B. D. Burks, holotype S (INHS); Se- 
rena, Indian Creek, 16-VI-1939, B. D. 
Burks, 1 6 (INHS); OHIO, Adams Co., 
Hills Fork-Eagle Creek, SR 125, W West 
Union, OH, 27-VI-1993, B. Armitage, UV 
trap, 37 S , 52 9 (OBS); OKLAHOMA, 
Latimer Co., lO-VI-1931, R. D. Bird, 3 6, 
3 $ (1 vial each sex, INHS). 

Discussion. — This species appears to be 



restricted to small streams in the Ohio and 
middle Mississippi River drainages. New 
state records are presented here for Ohio 
and Oklahoma. In addition to the states list- 
ed above, /. mazon has also been recorded 
from the Salt River drainage, Spencer Co., 
Kentucky (Resh 1975). 

Ithytrichia mexicana Harris and 

Contreras-Ramos 

(Figs. 3, 6, 9) 

Ithytrichia mexicana Harris and Contreras- 
Ramos 1989:176. 

Female description. — Length 3.5 mm. 20 
antennal segments. Brown in alcohol. Ster- 
nite VI with short acute ventro-mesal pro- 
cess. Sternite VII in ventral view subrect- 
angular. Sternite VIII in ventral view par- 
allel-sided, posterior margin with series of 
stout setae, each arising from a membra- 
nous tubercle; ventral sclerite with antero- 
lateral flaps, each bearing a series of short 
setae, narrowing to truncate posterior mar- 
gin, posterior margin with a short mesal 
process. Segment IX bullet-shaped in ven- 
tral view. Segment X button-like with a pair 
of short cerci. Apodemes slender, rod-like, 
extending from segment IX to VII, anterior 
apices straight, second pair of lateral apo- 
demes branching from mesal pair at ap- 
proximately the anterior margin of segment 
VIII. Spermathecal sclerite in ventral view 
with anterior portion forming a funnel- 
shaped collar constricted in middle; basal 
one-half sclerotized laterally with anterior 
apices acutely produced mesad, gradually 
tapering posteriorly to pointed, incurved 
apices, middle portion of sclerotized appa- 
ratus arrowhead-shaped; posterior one-half 
widest at base with lateral patches of mi- 
nute spines, narrowing in middle to tubular 
apex, middle portion with short mesal in- 
cision; anterior and posterior portions join- 
ing with lateral membranous lobes. 

Material examined.— USA: ARIZONA, 
Coconino Co., Oak Creek at Sterling Spring 
Fish Hatchery, U.S. Hwy 89A, elev. 1,829 
m, 4-5-VI-1993, S. Moulton and K. Alex- 



VOLUME 101. NUMBER 2 



237 




D 





B C 

Fig. 2. Ithytiichiu mazon, male genitalia. A, Left lateral. B. Dorsal. C, Ventral. D, Phallus. 



238 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




D 





Fig. 3. Ithytrichia mexicana, male genitalia. A, Left lateral. B, Dorsal. C, Ventral. D, Phallus. 



VOLUME 101, NUMBER 2 



239 




Figs. 4-6. Ithytrichia female genitalia, ventral. 4, /. clavata. 5, /. mazon. 6, /. mexicana. 



ander, 2 S (UM); NEW MEXICO, Sandov- 
al Co., Rito de los Frijoles @ Bandelier Na- 
tional Monument, 10.4 km S Los Alamos, 
elev. 1,839 m, 2-VIII-1994, L. E Carter, 1 
6 (INHS), same but 1-3-VIII-1997, J. Slu- 
sark and B. Richards, UV trap, 1 $ 
(NMNH). MEXICO: TAMAULIPAS, 
Municipio de Gomez Farias, Rio Frio @ 
headwaters. La Poza Azul, 6 km S Gomez 



Farias, 7-VIII-1988, A. Contreras and A. 
Moreno, blacklight, holotype 6 (NMNH). 
Discussion. — This species was originally 
described by Harris and Contreras-Ramos 
(1989) based on a single male collected 
from the headwaters of the Rio Frio, Ta- 
maulipas, Mexico. The Arizona record list- 
ed above was erroneously reported as a new 
Arizona state record for /. clavata by Moul- 



240 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 






8 



Figs. 7-9. Ithytrichia spermathecal sclerites, ventral. 7. /. cUivata. 8. /. nicizon. 9. I. mexicima. 



ton et al. (1994). Ithytrichia mexicana is re- 
ported herein from the United States for the 
first time and the species is now represented 
by a total of five specimens (4 males, 1 fe- 
male). On the basis of known collection rec- 
ords, it appears to have an affinity for small 
cold mountain streams in the southwestern 
United States and northern Mexico at about 
1,800 m in elevation. 

Undetermined Ithytrichia Material 

We examined several larvae and pupae 
in this study that could not be positively 
determined to species. However, specula- 
tion as to their probable identity is indicated 



in brackets for a few records based on dis- 
tributional information. 

Material examined. — ARIZONA, Gila 
Co., Christopher Creek, AZ 260, ca. 40 km 
NE Payson, elev. 1,792 m, 14- VII- 1985, A. 
R. Brigham, 2 larvae (INHS) [/. mexicana]; 
Christopher Creek, ?-?-1985, A. Brigham, 
8 larvae (INHS) [/. mexicana}; Lower Hor- 
ton Creek, 12-VIII-1937, Tazwell, 1 larva 
(INHS); ARKANSAS, Randolph Co., 
Jane's Creek, AR Hwy 90, S Ravenden 
Springs, III- 1985, S. R. Moulton, 1 larva 
(ASUMZ); MAINE, Washington Co., Nar- 
raguagus River, island, 6-VII-1973, T. Min- 
go, 2 pupae, 5 larvae (NMNH) [/. clavata]; 



VOLUME 101, NUMBER 2 



241 



TEXAS, Pecos River, Sheffield, 6-1-1976, 
J. Davis, 6 larvae (NMNH) [/. clavata]; 
WISCONSIN, Madison, Fox River, Lake 
Winnebago, ?-?-1954, K. M. Mackenthun, 
1 larva (INHS) [/. clavata]. 

Key to the North American 
Species of Ithytrichia 

1. Male (Figs. 1-3) 2 

- Female (Figs. 4-6) 4 

2. Inferior appendages in ventral view tapering 
posterad. apices rounded (Figs. IC, 3C) .... 3 

- Inferior appendages in ventral view rectangu- 
lar, apices truncate (Fig. 2C) /. mazon 

3. Posterolateral margins of tergum IX .sclerotized, 
hooked laterad (Fig. 3B); apex of subgenital plate 
emarginate (Fig. 3C) /. mexicana 

- Posterolateral margins of tergum IX rounded, 
not hooked (Fig. IB): apex of subgenital plate 
dome-shaped (Fig. IC) /. clavata 

4. Ventral sclerite of VIII gradually widening pos- 
teriorly (Figs. 4, 5); lateral sclerites of sper- 
mathecal sclerite less than one-half length of 
entire apparatus, ending anterolaterally in an- 
gled or rounded apices (Figs. 7, 8) 5 

- Ventral sclerite of VIII narrowing posteriorly 
(Fig. 6); lateral sclerites of spermathecal sclerite 
about one-half length of entire apparatus, an- 
terior apices curving inward to join arrowhead- 
shaped mesal process (Fig. 9) /. mexicana 

5. Ventral sclerite of VIII with concave posterior 
margin (Fig. 4); lateral processes of sperma- 
thecal sclerite short, anterolaterally angled out- 
ward (Fig. 7) /. clavata 

- Ventral sclerite of VIII with truncated posterior 
margin (Fig. 3): lateral processes of sperma- 
thecal sclerite elongate, rounded anteriorly 
(Fig. 8) /. Diazon 

Acknowledgments 

We thank Lisa Carter, Oliver Flint, Jr. 
(NMNH), George Harp (ASUMZ), David 
Houghton (UM), Vincent Lee (CAS), Boris 
Kondratieff (CSU), and Kathy Zieders 
(INHS) for loaning specimens from their 
collections. Brian Armitage (OBS) provid- 
ed us with light trap material from which 
the female of /. mazon was discovered. Bra- 
dy Richards (USGS) assisted JPS in the col- 
lection of the /. mexicana female. Brian Ja- 
cobs (Bandelier National Monument, NM) 
greatly facilitated the completion of this 



study by providing a scientific collector 
permit. Brian Armitage. Gregg Easley 
(USGS), Oliver Flint, Jr., Boris Kondratieff, 
Jon Raese (USGS), and John Sandberg 
(USGS) reviewed drafts of the manuscript. 

Literature Cited 

Eaton, A. E. 1873. On the Hydroptilidae, a family of 
the Trichoptera. Transactions of the Entomological 
Society of London 1873: 125-150. 

Fischer, F C. J. 1961. Trichopterorum Catalogiis Vol. 
II: Philopotamidae, Hydroptilidae, Stenopsychi- 
dae. iv + 190 pp. Amsterdam. 

Hanis, S. C. and A. Contreras-Ramos. 1989. Ithyrichia 
mexicana (Trichoptera: Hydroptilidae), a new spe- 
cies of caddisfly from Mexico. Entomological 
News 100: 176-178. 

Houghton, D. C. and K. W. Stewart. 1998. Seasonal 
flight periodicities of six microcaddisflies (Trichop- 
tera: Hydroptilidae, Glossosomatidae) in the Brazos 
River, Texas, with notes on larval biology and site 
records. Entomological News 109: 103-109. 

Marshall, J. E. 1979. A review of the genera of the 
Hydroptilidae (Trichoptera). Bulletin of the British 
Museum of Natural History (Entomology) 39: 
135-239. 

Morse, J. C. 1993. A checklist of the Trichoptera of 
North America, including Greenland and Mexico. 
Transactions of the American Entomological So- 
ciety 119: 47-93. 

Morton, K. J. 1905. New American Hydroptilidae. 
New York State Museum Bulletin 86: 63-85. 

Moulton, S. R., II, K. W. Stewart, and K. L. Young. 
1994. New records, distribution, and taxonomic 
status of some northern Arizona caddisflies (Tri- 
choptera). Entomological News 105: 164-174. 

Moulton, S. R., II and K. W. Stewart. 1996. Caddisflies 
(Trichoptera) of the Interior Highlands of North 
America. Memoirs of the American Entomologi- 
cal Institute Vol. 56, 313 pp. 

Resh, V. H. 1975. A distributional study of the cad- 
disflies of Kentucky. Transactions of the Kentucky 
Academy of Science 36: 6-16. 

Ross, H. H. 1944. The caddisflies, or Trichoptera, of 
Illinois. Bulletin of the Illinois Natural History 
Survey 23: 1-326. 

Wiggins, G. B. 1996. Larvae of the North American 
caddisfly genera (Trichoptera). 2nd Edition. Llni- 
versity of Toronto Press, Toronto. 457 pp. 

Wiggins, G. B. and C. R. Parker. 1997. Caddisflies 
(Trichoptera) of the Yukon, with analysis of the 
Beringian and holarctic species of North America, 
pp. 787-866. In Danks, H. V. and J. A. Downes, 
eds.. Insects of the Yukon. Biological Survey of 
Canada (Terrestrial Arthropods), Ottawa. 1034 pp. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999, pp. 242-262 

CHECKLIST AND HOST PLANTS OF THE TREEHOPPERS (HEMIPTERA: 
MEMBRACIDAE) OF NORTH CAROLINA 

Christopher H. Dietrich, Mark J. Rothschild, and Lewis L. Deitz 

(CHD) Center for Biodiversity, Illinois Natural History Survey, 607 East Peabody 
Drive, Champaign, IL 61820, U.S.A. (e-mail: dietrich@denrl.igis.uiuc.edu); (MJR) Mary- 
land Department of Agriculture, Plant Protection Section, 27722 Nanticoke Road, Unit 2, 
Salisbury, MD 21801, U.S.A.; (LLD) Department of Entomology, North Carolina State 
University, Raleigh, NC 27695-7613 (e-mail: ldietz@ppentl.ppath.ncsu.edu) 



Abstract. — Based on recent collecting and an examination of museum specimens, at least 
89 treehopper species (Hemiptera: Membracidae) occur in North Carolina, of which 26 
species represent new state records. The presence of 13 species previously recorded from 
North Carolina could not be verified based on available material. Three previous North 
Carolina records were found to be based on misidentifications. The known distribution (by 
county) and host plants in North Carolina are given for each species. Photographs of rep- 
resentative taxa and a host plant index are included. Stictocephala bisonia Kopp and Yonke 
is reinstated as a valid name (and not a junior synonym of Ceresa alta Walker). 

Key Words: Membracidae, treehopper, taxonomy, biogeography, insect-plant interactions 



The family Membracidae (Figs. 1-10) in- 
cludes more than 3,000 described treehopper 
species worldwide (McKamey 1998). About 
260 are known to occur in temperate North 
America. Many of these species are restrict- 
ed to the mixed hardwood forests and sa- 
vannas of the eastern United States, where 
they exploit a variety of woody and herba- 
ceous plants as hosts for oviposition, feed- 
ing, or both. Most North American treehop- 
per species are univoltine, solitary, and cryp- 
tic as both immatures and adults, and, hence 
are seldom noticed or collected. A few spe- 
cies, however, are multivoltine, gregarious 
(Figs. 7, 8), ant-mutualistic (Figs. 8, 10), or 
aposematic (Fig. 7), and are therefore some- 
what conspicuous. Three kinds of life cycles 
are common among North American tree- 
hoppers (Table 1). Many members of cate- 
gory III that feed and oviposit on oaks (Figs. 
5, 8) are usually found as adults for only a 
few weeks in May or June, depending on 
the location within the state. 



Records of North Carolina treehoppers 
were summarized by Metcalf (1915), Brim- 
ley (1938, 1942), Wray (1950, 1967), Met- 
calf and Wade (1965), and Kopp and Yonke 
(1973a-c, 1974: distribution maps). Kopp 
and Yonke's series provided keys to many 
species in eastern North America. Deitz et 
al. (1976), McGiffen and Neunzig (1985), 
and Hargrove (1986) gave further records 
of North Carolina treehoppers associated 
with soybeans, grapes, and black locust, re- 
spectively. 

The objectives of the present work were 
to document the species richness of North 
Carolina treehoppers, summarize the known 
distributions (Fig. 1 1 : county map) and host 
plant associations within the state, and pro- 
vide an up-to-date checklist following cur- 
rent nomenclature. Although host records 
for numerous species have been published 
(e.g., Funkhouser 1917; Ball 1931; Kopp 
and Yonke 1973a-c, 1974), the extent to 



VOLUME 101, NUMBER 2 



243 




Figs. 1-6. Representative treehoppers of North Carolina. 1, Microcentrus catyae. 2, Acutalis tartarea. 3, 
Micrutalis calva. 4. Campylenchia latipes. 5, Glossonotus univitlatiis. 6. Stictocephala niilitaris. 



which host associations vary geographical- 
ly is poorly documented and most pub- 
lished records have not been verified 
through rearing of immatures. 

Materials and Methods 

The species records below are based on 
specimens in the following collections: the 
North Carolina State University Insect Col- 
lection, Raleigh [NCSU: includes recent 
material from the authors]; Mark J. Roths- 



child Collection, % Maryland Department 
of Agriculture, Salisbury [MJRC]; Florida 
State Collection of Arthropods, Gainesville 
[FSCA]; North Carolina Department of Ag- 
riculture, Raleigh [NCDA]; and the Nation- 
al Museum of Natural History, Smithsonian 
Institution, Washington, D.C. [USNM]. C. 
S. Brimley's historically important material 
is housed at the collections of NCDA (spec- 
imens and card files) and NCSU (speci- 
mens). 



244 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 7-10. Representative treehoppers of Nortii Carolina (continued). 7, Platycoiis vittata, aggregation ot 
teneral adults. 8, Vcmdnzeu arqiuito. aggregation of adults and nymphs attended by ants. 9. Ophidernui evelyna: 
left, female (green), right, male (brown). 10, Entylici cahnata (left) and Piiblilia concavci (right, attended by 
ant). 



VOLUME 101. NUMBER 2 



245 



Table 1. Three major kinds of life cycles in treehoppers of eastern North America (modified from Kopp and 
Yonke 1973a). 



Calcgoiy: Taxa 



Overwintering 



Development 



Generations Per Year 



I: Polyglyptini. Platycotis 
vittata. probably Cam- 
pylenchia latipes, some 
Vandiiz.ea and some 
Ceresini 

II: Acutalini. most Ceresini. 
some Micnitalis 



III: Smiliini. Microcentnis 

spp., Enchenopa biiiota- 
ta complex, some Mi- 
crutalis 



adults overwinter in litter nymphs feed and develop 2 (most are bivoltine) 
on herbaceous or 
woody host plants 



eggs overwinter under nymphs feed and develop 1 (univoltine) 



bark in young twigs of 
woody hosts 



eggs overwinter under 
bark in young twigs of 
woody hosts 



on herbaceous host 
plants (many females 
require preoviposition 
period before laying 
eggs) 

nymphs feed and develop 1 (most are univoltine) 
on same woody hosts 
used for oviposition 



Records are based on adult specimens 
except as noted. Among species of Ceresi- 
ni, males are generally required for positive 
identification (Kopp and Yonke 1979); con- 
sequently, females without associated males 
often could not be identified. Each entry in- 
cludes a list of counties in North Carolina 
from which the species has been recorded, 
the seasonal distribution (earliest and latest 
calendar date of collection of adults), and 
North Carolina host records. A few speci- 
mens bore labels indicating a locality situ- 
ated on the border of two or three counties; 
these were considered to occur in all of the 
counties involved. 

Except as noted, only North Carolina 
host associations based on specimens ex- 
amined are reported here. Hosts marked 
with an asterisk (*) are those from which 
both nymphs and adults have been collect- 
ed. Other plants listed are those from which 
only adults have been collected, so some 
may not be true hosts. Botanical nomencla- 
ture follows Kartesz (1994), Liberty Hyde 
Bailey Hortorium (1976), and Radford et al. 
(1968). To conserve space, botanical com- 
mon names and the authors of plant scien- 
tific names are given only in the alphabet- 
ical host index. 

To facilitate comparisons with The In- 



sects of North Carolina (Brimley 1938, 
1942, Wray 1950, 1967), and The Treehop- 
pers of Missouri (Kopp and Yonke 1973a- 
c, 1974), names from those works that dif- 
fer from current nomenclature are given for 
each entry in square brackets (occasionally 
with other notes on synonymy). For addi- 
tional synonymy, see Fascicle 1, Membra- 
cidae, and its supplements, in the General 
Catalogue of the Hemiptera (Funkhouser 
1927, Metcalf and Wade 1965, McKamey 
1998) and the associated bibliographies 
(Metcalf and Wade 1963, Deitz and Kopp 
1987, Deitz 1989). 

For convenience, the checklist is ar- 
ranged alphabetically by genus and species. 
Table 2 summarizes the placement of the 
included genera into tribes and subfamilies. 

Results 

Recent collecting in North Carolina 
yielded numerous new state, county, and 
host records. Figure 1 1 indicates the num- 
ber of treehopper species recorded for each 
of North Carolina's 100 counties. The high- 
er species richness recorded near Raleigh 
(Wake County, 68 species), Asheville (Bun- 
combe County, 39 species), Boone (Watau- 
ga county, 30 species), and Charlotte 



246 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



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VOLUME 101, NUMBER 2 



247 



Table 2. Summary of the classification of North Carolina Membracidae (based on Deitz 1975, Kopp and 
Yonke 1979, and Deitz and Dietrich 1993). 

Subfamily Stegaspidinae: 

Tribe Microcentrini: Microcentnis Stal 1869 (Fig. 1). 
Subfamily Membracinae: 

Tribe Hoplophorionini: Playcotis Stal 1869 (Fig. 7). 

Tribe Membracini: Tylopelta Fowler 1894; Campylenchia Stal 1869 (Fig. 4); Enchenopa Amyot & Serville 
1843. 
Subfamily Smiliinae: 

Tribe Acutalini: Aciilalis Fairmaire 1846 (Fig. 2). 

Tribe Micrutalini: Micrittalis Fowler 1895 (Fig. 3). 

Tribe Ceresini: Hadrophallus Kopp and Yonke 1979; Stictolohus Metcalf 1916; Tortistilus Caldwell 1949; 

Stictocephala Stal 1869 (Fig. 6); Spissistilus Caldwell 1949. 
Tribe Amastrini: Vandiizea Coding 1892 (Fig. 8). 

Tribe Smiliini: Archasia Stal 1867; Carynota Fitch 1851; Glossonotus Butler 1877 (Fig. 5); Heliria Stal 1867; 
Telamona Fitch 1851; Thelia Amyot and Serville 1843; Atymna Stal 1867; Cyrtolobiis Coding 1892; Ophi- 
denna Fairmaire 1846 (Fig. 9); Smilia Germar 1833; Xantholobus Van Duzee 1908. 
Tribe Polyglyptini: Publilia Stal 1866 (Fig. 10, right); Entylia Germar 1833 (Fig. 10, left). 



(Mecklenburg County, 29 species) reflects 
greater collecting effort in those areas. 

With at least 89 species. North Carolina 
ranks third among the few states for which 
treehopper checklists have been published. 
New York ranks first (100 currently recog- 
nized species: Leonard 1928) and Ohio sec- 
ond (93 currently recognized species, in- 
cluding two presumed present based on rec- 
ords in neighboring states: Osbom 1940), 
but dubious identifications may have inflat- 
ed the accuracy of counts for those states. 
Among treehoppers, males, females, or 
both may be polymorphic with respect to 
pronotal shape and coloration (Figs. 7, 9). 
Frequently, previous workers incorrectly 
identified a single polymorphic species as 
two or more distinct species. 

Thirteen species previously reported 
from North Carolina whose occurrence in 
the state we could not verify and three pre- 



Table 3. Comparison of treehopper diversity in the 
world, the Nearctic Region, and North Carolina (based 
on McKamey 1998 and the present work). 



Geographic 
Area 



Species 



Genera 



Suh- 
familit 



World 

Nearctic Region 
North Carolina 



3.177 
258 



397 
62 
26 



49 

13 

9 



12 
6 

3 



viously misidentified species — Hadrophal- 
lus constans (Walker), Spissistilus unifor- 
mis (Fairmaire), and Telamona concava 
Fitch, are included below in square brack- 
ets. 

Five additional species reported from ad- 
jacent states may eventually be found in 
North Carolina, but are not included in the 
checklist: Palonica pyramidata (Uhler), 
Stictocephala albescens (Van Duzee), and 
Thelia uhleri Stal, all reported from Virgin- 
ia (Kopp and Yonke 1973b, 1974: distri- 
bution maps); Helonica excelsa (Fairmaire), 
reported from South Carolina (Kopp and 
Yonke 1974: map); and Telamona compac- 
ta Ball (specimen [USNM] from Rocky 
Bottom, South Carolina [<10 miles from 
Transylvania County, North Carolina]). 

Table 3 shows the species richness of 
North Carolina's treehopper fauna com- 
pared to the Nearctic Region and the world. 
Especially well represented in North Caro- 
lina are the genera Cyrtolobus (21 species), 
Telamona (10), Ophiderma (7), and Heliria 
(6). All species recorded are native to North 
Carolina and nearly all are endemic to the 
eastern U.S. 

North Carolina treehoppers have been 
collected on at least 45 plant genera in 19 
families, including 12 genera of Asteraceae, 



248 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



9 of Fabaceae, and 3 of Fagaceae. More- 
over, 39 species (43 percent of the state's 
membracid fauna) may be found on oaks 
(Fagaceae), with 16 species of Quercus Ust- 
ed below as hosts. 

Checklist of North Carolina 
Membracidae 

Acutalis tartarea (Say 1830) [in part as 
Acutalis tartarea var. semicrema (Say 
1830) in Brimley 1938]. Fig. 2. Counties: 
Alamance, Alexander, Anson, Ashe, 
Beaufort, Bladen, Buncombe, Burke, Ca- 
barrus, Caldwell, Carteret, Chatham, Da- 
vidson, Duplin, Edgecombe, Forsyth, 
Granville, Haywood, Henderson, Hyde, 
Jackson, Johnston, Mecklenburg, Nash, 
Onslow, Orange, Pamlico, Pasquotank, 
Person, Polk, Rutherford, Stanly, Surry, 
Tyrrell, Wake, Warren, Washington, Wa- 
tauga, Wayne, Wilkes, Wilson. Seasonal 
distribution: 24 May-23 November. Host 
associations: Ambrosia artemisiifolia, 
Arimdinaria sp., Bidens coronata, Eu- 
patorium capiUifolium, Helianthus sp., 
Solidago sp. 

Archasia auriculata (Fitch 1851) [as A. gal- 
eata (Fabricius 1803), a preoccupied 
name, in Brimley 1938]. Counties: Blad- 
en, Buncombe, Columbus, Moore, Wake. 
Seasonal distribution: 9 May-27 August. 
Host associations: Quercus nigra, Q. ve- 
lutina"^. 

Archasia belfragei Stal 1869. Counties: 
Bladen, Buncombe, Columbus, Hender- 
son, Macon, Wake. Seasonal distribution: 
12 May-1 1 September. Host associations: 
Quercus alba*, Q. phellos. Notes: Brim- 
ley's (1938) record of A. belfragei from 
Southern Pines (Moore County, June), 
actually represents a specimen (NCDA) 
of A. auriculata. 

Atymna castaneae (Fitch 1851) [as Cyrto- 
lobus castaneus (Fitch) in Brimley 1938]. 
Counties: Ashe, Bladen, Buncombe, 
Burke, Graham, Haywood, Jackson, 
Sampson, Surry, Swain, Watauga, Yan- 
cey. Seasonal distribution: 23 May-31 



August. Host associations: Castanea den- 
tata*, C. pumila*. 

[Atymna inornata (Say 1830)]. Records of 
this species in North Carolina (Metcalf 
1915, Van Duzee 1917, Brimley 1938 [as 
Cyrtolobus inornata]) were not verified 
by the present authors. 

Atymna querci (Fitch 1851). Counties: Al- 
leghany, Ashe, Buncombe, Cabarrus, 
Cleveland, Duplin, Forsyth, Haywood, 
Macon, Martin, Nash, Orange, Vance, 
Wake, Warren, Washington, Watauga, 
Wayne, Yadkin. Seasonal distribution: 14 
April-8 September. Host associations: 
Quercus alba*, Q. stellata*, Vitis rotun- 
difolia. 

Campylenchia latipes (Say 1824). Fig. 4. 
Counties: Alamance, Alexander, Allegh- 
any, Ashe, Avery, Bladen, Brunswick, 
Buncombe, Burke, Cabarrus, Caldwell, 
Chatham, Cleveland, Davidson, Davie, 
Duplin, Durham, Forsyth, Graham, Gran- 
ville, Guilford, Halifax, Haywood, John- 
ston, Jones, Macon, Madison, Martin, 
McDowell, Mecklenburg, Mitchell, 
Montgomery, Moore, New Hanover, 
Onslow, Orange, Pender, Polk, Ruther- 
ford, Sampson, Scotland, Stanly, Swain, 
Transylvania, Wake, WaiTen, Washing- 
ton, Watauga, Wayne, Wilkes, Yadkin. 
Seasonal distribution: [? 4 May (NCSU)], 
3 June— 21 November. Host associations: 
Medicago sativa, Solidago sp. 

Carynota marmorata (Say 1830). Counties: 
Haywood, Henderson, Jackson, Moore, 
Stanly, Wake, Watauga. Seasonal distri- 
bution: 6 June-30 July. Host associa- 
tions: Betula sp. 

Carynota mera (Say 1830). Counties: Blad- 
en, Currituck, Hyde, Mecklenburg, New 
Hanover, Pitt, Wake. Seasonal distribu- 
tion: [18 May, nymphs], 21 May-late Oc- 
tober. Host associations: Carya illino- 
inensis, C. sp*. 

Cyrtolobus arcuatus (Emmons 1854). 
Counties: Greene, Harnett, Hertford, 
Wake. Seasonal distribution: 24 April— 12 
May. Host associations: Quercus falca- 
ta*, Q. phellos. 



VOLUME 101. NUMBER 2 



249 



Cyrtolobus auroreus Woodruff 1924. NEW 
STATE RECORD. Counties: Ashe, Du- 
plin, Guilford, Macon, Mecklenburg, 
Wake, Wayne. Seasonal distribution: 23 
April-19-20 June. Host associations: 
Qiiercus alba*, Q. prinus. 

Cyrtolobus celsus Van Duzee 1917 [as C. 
celsis (sic) in Brimley 1938]. Counties: 
Moore. Seasonal distribution: 22 May. 
Host associations: no data for North Car- 
olina. 

Cyrtolobus clarus Woodruff 1924. NEW 
STATE RECORD. Counties: Wake. Sea- 
sonal distribution: 4-25 May. Host as- 
sociations: Quercus palustris*, Q. phel- 
los*. 

[Cyrtolobus discoidalis (Emmons 1854)]. 
Brimley 's (1938) record of this species in 
North Carolina (Balsam, Jackson Coun- 
ty) could not be verified, but may be the 
basis for Kopp and Yonke's (1973c: map) 
record. 

Cyrtolobus dixiamis Woodruff 1924. NEW 
STATE RECORD. Counties: Cabarrus, 
Duplin, Franklin, Wake. Seasonal distri- 
bution: 26 April-26 May. Host associa- 
tions: Quercus alba, Q. falcata, Q. pal- 
ustris, Q. stellata*. 

Cyrtolobus fene stratus (Fitch 1851). Coun- 
ties: Bladen, Dare, Edgecombe, Forsyth, 
Greene, Hertford, Hoke, Johnston, Le- 
noir, Nash, Pasquotank, Robeson, Rock- 
ingham, Wake, Washington, Wilson. Sea- 
sonal distribution: [15 April, nymph], 24 
April- 19 May. Host associations: Quer- 
cus falcata*, Q. laevis, Q. margarettiae, 
Q. marilandica*, Q. nigra*, Q. palustris, 
Q. phellos*, Q. rubra*, Q. velutina, Q. 
virginiana. Notes: Brimley 's (1938) June 
record of C fenestratus from Blowing 
Rock (Watauga County) was not verified 
by the present authors, while his July rec- 
ord from Blowing Rock may represent a 
specimen (NCDA) of C puritanus 
Woodruff. 

Cyrtolobus flavolatus Woodruff 1924. 
NEW STATE RECORD. Counties: Bun- 
combe, Vance, Wake. Seasonal distribu- 



tion: 18 May- 18 June. Host associations: 
no data for North Carolina. 

Cyrtolobus fuliginosus (Emmons 1854). 
Counties: Cabarrus, Franklin, Greene, 
Harnett, Hertford, Johnston, Lenoir, Mar- 
tin, Mecklenburg, Pasquotank, Randolph, 
Wake, Warren, Washington, Wayne. Sea- 
sonal distribution: 23 April-26 May. 
Host associations: Quercus coccinea, Q. 
falcata*, Q. palustris, Q. phellos, Q. stel- 
lata, Q. velutina*. 

Cyrtolobus funkhouseri Woodruff 1924. 
NEW STATE RECORD. Counties: 
Mecklenburg, Stokes, Wake, Wilkes, 
Yadkin. Seasonal distribution: 13 May- 
1 1 June. Host associations: Quercus pal- 
ustris. 

Cyrtolobus fuscipennis Van Duzee 1908. 
Counties: Ashe, Buncombe, Burke, Jack- 
son, Macon, Mecklenburg, Wake. Sea- 
sonal distribution: 24 April-22 July. Host 
associations: Quercus alba*, Q. nigra. 

Cyrtolobus g risen s Van Duzee 1908. NEW 
STATE RECORD. Counties: Ashe, 
Burke, Montgomery, Vance, Wake. Sea- 
sonal distribution: 29 April-6 July. Host 
associations: Quercus alba, Q. stellata. 

Cyrtolobus inermis (Emmons 1854). NEW 
STATE RECORD. Counties: Bladen, 
Hoke, Rutherford, Wake. Seasonal distri- 
bution: 3 May-10 June. Host associa- 
tions: Quercus falcata* , Q. marilandica* , 
Q. stellata. 

Cyrtolobus maculifrontis (Emmons 1854). 
Counties: Ashe, Bladen, Duplin, Gran- 
ville, Vance, Wake, Watauga, Wayne. 
Seasonal distribution: [? March (NCSU)], 
23 April-27 June. Host associations: 
Quercus alba*, Q. falcata. 

Cyrtolobus ovatus Van Duzee 1908. Coun- 
ties: Bladen, Hoke, Johnston, Moore, 
Richmond, Sampson, Wake. Seasonal 
distribution: 7 May-28 June. Host asso- 
ciations: Quercus laevis*, Q. marilandi- 
ca*. 

Cyrtolobus pallidifrontis (Emmons 1854). 
NEW STATE RECORD. Counties: Bun- 
combe, Currituck, Wake, WaiTcn, Watau- 
ga. Seasonal distribution: 29 April-26 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



July. Host associations: Quercus alba, Q. 
stellata*. 

Cyrtolobus parx'ulus Woodruff 1924. NEW 
STATE RECORD. Counties: Bladen, 
Moore, Richmond. Seasonal distribution: 
23 May-17 June. Host associations: no 
data for North Carolina. Notes: Hosts re- 
ported elsewhere include two species of 
Quercus (Kopp and Yonke 1973c). 

Cyrtolobus pulchellus Woodruff 1924. 
NEW STATE RECORD. Counties: Ashe. 
Seasonal distribution: 19-20 June. Host 
associations: Quercus rubra. 

Cyrtolobus puritanus Woodruff 1924. 
Counties: Buncombe, Wake, Watauga, 
Yancey [as "Black Mountains," which is 
on the Buncombe- Yancey border 
(USNM)]. Seasonal distribution: 26 
May-20 July. Host associations: no data 
for North Carolina. Notes: Brimley's 
(1938) record of C. puritanus from Lake 
Toxoway (Transylvania Co.) was not ver- 
ified by the present authors. Hosts re- 
ported elsewhere include five species of 
Quercus (Kopp and Yonke 1973c). 

[Cyrtolobus sculptus (Fairmaire 1846)]. Rec- 
ords of C sculptus in North Carolina 
(Coding 1893, Van Duzee 1917, Brimley 
1938) were not verified by the present au- 
thors. 

Cyrtolobus togatus Woodruff 1924. NEW 
STATE RECORD. Counties: Buncombe, 
Cabarrus, Durham, Franklin, Harnett, 
Mecklenburg, Nash, Rockingham, Vance, 
Wake, Yadkin. Seasonal distribution: 23 
April-23-30 June. Host associations: 
Quercus nigra*, Q. phellos*, Q. stellata. 

Cyrtolobus tuberosus (Fairmaire 1846). 
Counties: Bladen, Cabarrus, Camden, 
Caswell, Columbus, Craven, Franklin, 
Hertford, Hoke, Lenoir, Mecklenburg, 
Montgomery, Nash, Northampton, Robe- 
son, Rockingham, Wake, Warren, Wayne, 
Wilson. Seasonal distribution: 16 April- 
1 June. Host associations: Quercus alba*, 
Q. margarettiae, Q. marilandica, Q. ni- 
gra, Q. prinus, Q. rubra, Q. stellata, Q. 
virginiana. 

Cyrtolobus vau (Say 1830). Counties: Al- 



leghany, Ashe, Avery, Buncombe, Ca- 
swell, Columbus, Duplin, Franklin, Hay- 
wood, Henderson, Jackson, Macon, 
Mecklenburg, Moore, Rockingham, Tran- 
sylvania, Vance, Wake, Yancey. Seasonal 
distribution: late March-5 September. 
Host associations: Quercus alba*, Q. pri- 
nus, Q. stellata. 
Enchenopa binotata (Say 1824) complex 
(see Notes, below, for discussion of the 
complex). Counties: Alamance, Allegh- 
any, Ashe, Buncombe, Carteret, Chat- 
ham, Gates, Graham, Guilford, Hay- 
wood, Iredell, Macon, Madison, McDow- 
ell, Moore, New Hanover, Richmond, 
Sampson, Stanly, Transylvania, Wake, 
Yancey. Seasonal distribution: [1 May, 
nymph], 15 May-3 October. Host asso- 
ciations: Carya sp., Cercis canadensis* , 
Juglans nigra*, Liriodendron tulipifera, 
Robinia pseudoacacia* , Viburnum prun- 
ifolium*. Notes: The Enchenopa binotata 
complex is thought to include nine bio- 
logically distinct North American spe- 
cies, each of which is associated with a 
different genus or species of deciduous 
woody host plant: (1) Carya spp., (2) Ce- 
lastris scandens, (3) Cercis canadensis, 
(4) Juglans cinerea, (5) J. nigra, (6) Lir- 
iodendron tulipifera, (7) Ptelea trifoliata, 
(8) Robinia pseudoacacia, and (9) Vibur- 
num spp. (Pratt and Wood 1992, 1993). 
Three published names are currently 
available for species within this complex, 
but the corresponding original descrip- 
tions lack host plant data as well as mor- 
phological criteria useful for distinguish- 
ing either the nymphs or adults from oth- 
er species in the complex (Pratt and 
Wood 1992). Pratt and Wood (1992) de- 
scribed the fifth instar nymphs of species 
in the complex and provided a key for 
their identification, but did not attempt to 
resolve the nomenclatural problems. 

Based on host data, specimens examined 
from North Carolina appear to represent 
five of the nine species in the complex: 



VOLUME 101, NUMBER 



251 



Cercis canadensis"^: Buncombe and Wake 

Counties (10 June-5 July). 
Jiiglans nigra*: Ashe and Sampson County 

([23 May, nymph] 5 July). 
Liriodendron tulipifera: Chatham County 

(2 July). 
Robinia pseudoacacia*: Macon County (5 

August). 
Viburnum prunifolium*: Wake County (27 

May). 

Entylia carinata (Forster 1771) [in part as 
E. concisa Walker 1851, and as E. sinu- 
ata (Fabricius 1798) in Brimley 1938; as 
E. bactriana Germar 1835, in Kopp and 
Yonke 1973b; E. carinata (Forster) in 
Remes-Lenicov 1973]. Fig. 10 (left). 
Counties: Alamance, Alexander, Allegh- 
any, Anson, Ashe, Avery, Beaufort, Ber- 
tie, Bladen, Buncombe, Burke, Cabarrus, 
Caldwell, Camden, Catawba, Chatham, 
Cherokee, Clay, Cleveland, Cumberland, 
Dare, Davidson, Davie, Duplin, Durham, 
Edgecombe, Gaston, Gates, Graham, 
Granville, Harnett, Haywood, Henderson, 
Hertford, Hoke, Hyde, Iredell, Jackson, 
Johnston, Lincoln, Macon, Madison, Mc- 
Dowell, Mecklenburg, Montgomery, 
Moore, Nash, New Hanover, Onslow, 
Orange, Pasquotank, Perquimans, Pitt, 
Polk, Rockingham, Sampson, Scotland, 
Stanly, Surry, Swain, Transylvania, 
Vance, Wake, Warren, Washington, Wa- 
tauga, Wayne, Yadkin, Yancey. Seasonal 
distribution: 13 March-19 December. 
Host associations: Ambrosia artemisiifol- 
ia*. Ambrosia sp.. Aster sp., Bidens bi- 
pinnata, B. coronata, B. sp., Conyza can- 
adensis (as Erigeron canadensis). Dahlia 
sp., Erechtites hieraciifolia*, Erigeron 
sp., Eupatorium capillifolium, E. pilos- 
um, E. sp.. Glycine max, Helianthus an- 
nuus, H. tuberosus*, H. sp., Quercus pai- 
ns tris, Silphium sp.. Solan um tuberosum, 
Solidago sp., Verbesina alternifolia (as 
Actinomeris alternifolia), Vitis rotundi- 
folia, V. sp. Notes: Deitz et al. (1976) re- 
ported an Entylia from Glycine max in 
Columbus Co., however, the voucher ma- 



terial could not be located for the present 
study. Also, Brimley 's (1938) material 
from Hendersonville (Henderson County) 
and Willard (Pender County) could not 
be located and his specimen (NCDA) 
from "Spruce" is actually labelled "Sun- 
burst" (Haywood County). 

Glossonotus acuminatus (Fabricius 1775). 
Counties: Duplin, Wake. Seasonal distri- 
bution: 6 May-14 June. Host associa- 
tions: Quercus falcata*. 

Glossonotus turriculatus (Emmons 1854). 
Counties: Buncombe- Yancey [as "Black 
Mountains," which is on the border of 
these counties (USNM)]. Seasonal distri- 
bution: 15-20 June. Host associations: no 
data for North Carolina. Notes: Hosts re- 
ported elsewhere include Crataegus and 
Quercus (Kopp and Yonke 1974). 

Glossonotus univittatus (Harris 1841). Fig. 
5. Counties: Bladen, Burke, Mecklen- 
burg, Rockingham. Seasonal distribution: 
16 May-4 August. Host associations: 
Quercus alba, Q. rubra. 

Hadrophallus borealis (Fairmaire 1 846) [as 
Ceresa borealis Fairmaire in Brimley 
1938; as Spissistilus borealis (Fairmaire) 
in Kopp and Yonke 1973b]. Counties: 
Buncombe, Stanly, Swain, Wake, Watau- 
ga. Seasonal distribution: 3 June-29 Au- 
gust. Host associations: no data for North 
Carolina. Notes: One female specimen 
from Waynes ville, Haywood County, 14 
Sept., probably H. borealis-was formerly 
misidentified as Ceresa constans (Walk- 
er) by Z.R Metcalf (NCDA). Hosts re- 
ported elsewhere include species in sev- 
eral plant families (Kopp and Yonke 
1973b). 

[Hadrophallus constans (Walker 1851), 
misidentification]. Brimley 's (1938) rec- 
ords of Ceresa constans (Walker) refer to 
H. borealis (Raleigh [Wake County], 
Waynesville [Haywood County]), Sticto- 
cephala militaris (Havelock), or S. brev- 
ity I us (Newton); Kopp and Yonke 's 
(1973b: map) North Carolina record of 
this species (as S. constans) was probably 
based on Brimley's publication. 



252 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Heliria cornutula Ball 1925. NEW STATE 
RECORD. Counties: Bladen, Mecklen- 
burg, Randolph, Wake. Seasonal distri- 
bution: 5 July- 15 November. Host asso- 
ciations: no data for North Carolina. 

Heliria cristata (Fairmaire 1846). Counties: 
[as "Eastern N.C." on data label 
(NCSU)]. Seasonal distribution: mid 
July. Host associations: no data for North 
Carolina. Notes: Elsewhere, Kopp and 
Yonke (1974) reported Quercus macro- 
carpa Michx. 

Heliria gemma Ball 1925. NEW STATE 
RECORD. Counties: Graham. Seasonal 
distribution: 1 September. Host associa- 
tions: no data for North Carolina. 

Heliria gibberata Ball 1925. NEW STATE 
RECORD. Counties: Burke, Wake. Sea- 
sonal distribution: 23 May-late June. 
Host associations: no data for North Car- 
olina. Notes: Elsewhere, Ball (1931) re- 
corded Celtis occideiitalis L. as a host. 

[Heliria mexicana Stal 1869]. Records of 
H. mexicana in North Carolina (Ball 
1931, Brimley 1938) were not verified by 
the present authors. 

Heliria molaris (Butler 1877). Counties: 
Wake, Watauga. Seasonal distribution: 
August. Host associations: no data for 
North Carolina. Notes: Elsewhere, Kopp 
and Yonke (1974) reported Quercus and 
Populus as hosts. 

Heliria scalaris (Fairmaire 1846). Counties: 
Wake. Seasonal distribution: 21 June. 
Host associations: no data for North Car- 
olina. Notes: Elsewhere, Kopp and Yon- 
ke (1974) reported Crateagus as a host. 

Microcentrus caryae (Fitch 1851). Fig. 1. 
Counties: Buncombe, Forsyth, Haywood, 
Martin, Pitt, Sampson, Wake. Seasonal 
distribution: [7 May, nymph], 25 June-4 
December. Host associations: Carya illi- 
noinensis*, Carya sp. 

Microcentrus perditus (Amyot and Serville 
1843). Counties: Moore, Stokes, Wake. 
Seasonal distribution: 20 May-20 Octo- 
ber. Host associations: Carya illinoinen- 
sis. 

Micrutalis calva (Say 1830) [in part as Mi- 



cru talis calva var. illinoiensis (Coding 
1893) in Brimley 1938]. Fig. 3. Counties: 
Alamance, Alexander, Alleghany, Avery, 
Bertie, Bladen, Brunswick, Buncombe, 
Cabarrus, Caswell, Chatham, Cherokee, 
Columbus, Cumberland, Dare, Davidson, 
Davie, Duplin, Durham, Forsyth, Gra- 
ham, Granville, Guilford, Haywood, 
Hoke, Jackson, Johnston, Lincoln, Meck- 
lenburg, Mitchell, Moore, New Hanover, 
Onslow, Person, Richmond, Rutherford, 
Scotland, Stanly, Surry, Swain, Wake, 
Warren, Wayne, Wilkes. Seasonal distri- 
bution: late April-2 November. Host as- 
sociations: Conyza canadensis, Erigeron 
annuus, Gleditsia triacanthos*. Glycine 
max, Robinia pseudoacacia, Salix nigra, 
Solidago sp., Vitis rotundifolia, V. sp. 
'French hybrid'. Notes: Brimley 's (1938) 
records from Tin City and Willard (Pen- 
der County) were not verified by the pre- 
sent authors. 

Micrutalis dorsalis (Fitch 1851). Counties: 
Alleghany, Burke, Caldwell, Haywood, 
Swain, Watauga, Yancey. Seasonal distri- 
bution: 11 July-21 August. Host associ- 
ations: no data for North Carolina. 

Micrutalis malleifera Fowler 1895. NEW 
STATE RECORD. Counties: Dare. Sea- 
sonal distribution: 14 June. Host associ- 
ations: Physalis sp. Notes: Mead (1986) 
reviewed information on this treehopper, 
the only known vector of pseudo-curly 
top virus (a minor disease of tomatoes). 

Ophiderma definita Woodruff 1919. NEW 
STATE RECORD. Counties: Alamance, 
Beaufort, Bladen, Buncombe, Cabarrus, 
Edgecombe, Franklin, Harnett, Mecklen- 
burg, Nash, Northampton, Pitt, Polk, 
Randolph, Wake, Warren, Watauga. Sea- 
sonal distribution: 23 April-16 July, [11 
Sept. at UV-Hght (NCSU)]. Host associ- 
ations: Fagus grandifolia, Quercus fal- 
cata, Q. laurifolia, Q. nigra*, Q. palus- 
tris, Q. p he 1 1 OS*. 

Ophiderma evelyna Woodruff 1919. Fig. 9. 
Counties: Alamance, Bladen, Cabarrus, 
Greene, Harnett, Johnston, Lenoir, Meck- 
lenburg, Montgomery, Nash, Pitt, Ran- 



VOLUME 101, NUMBER 2 



253 



dolph. Wake, Washington, Wayne, Wil- 
son. Seasonal distribution: late March- 
23—30 June. Host associations: Quercus 
alba, Q.falcata*, Q. marilandica*, Q. ni- 
gra, Q. palustris, Q. phellos*. 

Ophiderma flava Coding 1893. Counties: 
Ashe, Buncombe-Haywood [as "Mt. Pis- 
gah," which is on the border of these 
counties], Macon, Watauga. Seasonal dis- 
tribution: 1 June-31 August. Host asso- 
ciations: Quercus alba*, Q. rubra. 

Ophiderma flavicephala Coding 1893. 
Counties: Alamance, Beaufort, Bun- 
combe, Cabarrus, Durham, Franklin, 
Hoke, Lenoir, Nash, Pitt, Rutherford, 
Wake, Wayne, Wilkes, Wilson, Yadkin. 
Seasonal distribution: 23 April-18 June. 
Host associations: Quercus alba, Q. coc- 
cinea, Q. falcata*, Q. nigra*, Q. palus- 
tris, Q. phellos*, Q. rubra var. ambigua 
(as borealis), Q. stellata. 

Ophiderma grisea Woodruff 1919. Coun- 
ties: Buncombe, Yancey [as "Valley of 
Black Mountains," which is on the Bun- 
combe-Yancey border (USNM)]. Sea- 
sonal distribution: 18 June-9 August. 
Host associations: no data for North Car- 
olina. Notes: Elsewhere, Kopp and Yon- 
ke (1973c) reported four species of Quer- 
cus as hosts. 

Ophiderma pubescens (Emmons 1854) [in 
part as Ophiderma flaviguttula Coding 
1893, and as Ophiderma pubescens var. 
australis Woodruff 1919, in Brimley 
1938]. Counties: Ashe, Hoke, Jackson, 
Johnston, Moore, Scotland, Wake. Sea- 
sonal distribution: 24 April-26 July. Host 
associations: Quercus falcata*, Q. mari- 
landica, Q. stellata. Notes: Brimley 's 
(1938) record of O. flaviguttata [misiden- 
tification] from Raleigh [Wake County], 
May, was based on a specimen of Cyr- 
tolobus flavolatus (NCDA). 

Ophiderma salamandra Fairmaire 1846. 
Counties: Ashe, Burke, Greene, Harnett, 
Johnston, Lee, Nash, Pasquotank, Wake, 
Warren, Watauga, Wayne, Wilson. Sea- 
sonal distribution: 18 April-31 August. 
Host associations: Quercus falcata, Q. ni- 



gra, Q. palustris, Q. phellos*, Q. rubra, 
Q. stellata, Q. velutina*. 

Platy'cotis vittata (Fabricius 1803) [as Pla- 
tycotis vittata var. quadrivittata (Say 
1830) in Brimley 1938]. Fig. 7. Counties: 
Alamance, Alexander, Alleghany, Anson, 
Ashe, Avery, Bladen, Buncombe, Burke, 
Cabarrus, Caswell, Catawba, Chatham, 
Cherokee, Chowan, Clay, Columbus, 
Craven, Cumberland, Dare, Durham, 
Forsyth, Franklin, Gaston, Graham, 
Granville, Guilford, Halifax, Haywood, 
Henderson, Hertford, Hoke, Iredell, Jack- 
son, Lee, Lenoir, Lincoln, Macon, Mar- 
tin, Mecklenburg, Montgomery, Moore, 
Nash, New Hanover, Northampton, Ons- 
low, Pasquotank, Pitt, Randolph, Robe- 
son, Rockingham, Sampson, Stanly, 
Stokes, Surry, Swain, Vance, Wake, War- 
ren, Watauga, Wilkes, Wilson, Yadkin, 
Yancey. Seasonal distribution: 16 Janu- 
ary-29 December. Host associations: Bet- 
ula sp.*, Castanea dentata*, Fagus sp.*, 
Quercus alba*, Q. falcata*, Q. incana*, 
Q. laevis, Q. margarettiae, Q. nigra*, Q. 
palustris* , Q. phellos, Q. rubra*, Q. rub- 
ra var. ambigua, Q. stellata* , Q. velutina, 
Q. virginiana*. 

Publilia concava (Say 1824). Fig. 10 
(right). Counties: Ashe, Avery, Bun- 
combe, Burke, Cherokee, Graham, Hay- 
wood, Henderson, Jackson, Macon, Mad- 
ison, McDowell, Mitchell, Swain, Watau- 
ga, Wilkes, Yadkin. Seasonal distribu- 
tion: late May-7 October. Host 
associations: Ambrosia artemisiifolia* , A. 
sp.*, Eupatorium sp.*, Helianthus sp.*, 
Solidago sp.* Notes: Brimley's (1938) 
specimen from "Spruce" is actually la- 
belled "Sunburst" (Haywood County: 
NCDA). 

Publilia reticulata Van Duzee 1908. Coun- 
ties: Ashe, Avery, Buncombe, Caldwell, 
Haywood, Henderson, Jackson, Transyl- 
vania, Wake, Watauga, Yancey (as 
"Black Mountains," which is on the 
Buncombe-Yancey border (USNM)]. 
Seasonal distribution: 29 April-4 Octo- 
ber, host associations: no data for North 



254 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Carolina. Notes: Hosts reported else- 
where include members of the Asteracae 
(Kopp and Yonke 1973b). 

Smilia camelus (Fabricius 1803). Counties: 
Burke, Greene, Hoke, Mecklenburg, 
Moore, Pender, Robeson, Sampson, 
Wake, Watauga. Seasonal distribution: 24 
April-25 September. Host associations: 
Quercus falcata*, Q. laevis*, Q. nigra*, 
Q. stellata. Notes: Brimley's (1938) re- 
cord of this species from Linville Falls 
[Caldwell County] was not verified by 
the present authors. 

Smilia fasciata Amyot and Serville 1843. 
NEW STATE RECORD. Counties: 
Alamance, Beaufort, Cabarrus, Chatham, 
Craven, Davidson, Franklin, Gaston, 
Harnett, Johnston, Lenoir, Mecklenburg, 
Moore, Nash, Orange, Pasquotank, Pen- 
der, Rockingham, Wake, Warren, Wayne, 
Wilson. Seasonal distribution: 2 April-5 
September. Host associations: Carya il- 
linoinensis, Quercus coccinea, Q. falca- 
m*, Q. nigra*, Q. palustris*, Q. phellos*, 
Q. stellata*, Q. velutina. 

[Spissistilus femoratiis (Fairmaire 1846)]. 
Records of this species [as Ceresa an- 
gulata (Walker 1851)] in North Carolina 
(Van Duzee 1917, Brimley 1938) were 
not verified by the present authors. 

Spissistilus festinus (Say 1830) [as Sticto- 
cephala festina (Say) in Brimley 1938]. 
Counties: Alexander, Anson, Beaufort, 
Bertie, Bladen, Brunswick, Buncombe, 
Cabarrus, Caldwell, Camden, Carteret, 
Catawba, Chatham, Chowan, Cleveland, 
Columbus, Craven, Cumberland, Curri- 
tuck, Dare, Davie, Duplin, Durham, Edge- 
combe, Franklin, Gaston, Gates, Gran- 
ville, Greene, Halifax, Harnett, Hay- 
wood, Henderson, Hoke, Iredell, Jackson, 
Johnston, Jones, Lee, Lincoln, Madison, 
Martin, Mecklenburg, Mitchell, Mont- 
gomery, Moore, Nash, New Hanover, 
Onslow, Orange, Pamlico, Pasquotank, 
Pender, Perquimans, Polk, Randolph, 
Robeson, Rockingham, Rowan, Samp- 
son, Scotland, Stanly, Swain, Transylva- 
nia, Tyrrell, Union, Vance, Wake, War- 



ren, Washington, Wayne, Wilkes, Wilson. 
Seasonal distribution: 19 February- 19 
December Host associations: Arachis hy- 
pogaea*. Aster ericoides. Glycine max*, 
Helianthus sp., Lespedeza sp.*, Medica- 
go sativa*, Phaseolits vulgaris*, Sarra- 
cenia flava, Solidago sp. Notes: This spe- 
cies is commonly known as the "three- 
cornered alfalfa hopper." Hargrove's 
(1986) record of S. festinus from Cow- 
eeta, Macon County (on Robinia pseu- 
doacacia), was not verified by the present 
authors. 

[Spissistilus rotundata Stal 1869]. Brim- 
ley's (1938) records of Stictocephala ro- 
tundata Stal in North Carolina (Raleigh 
[Wake County] and Southern Pines 
[Moore County]) were not verified by the 
present authors. 

[Spissistilus uniformis (Fairmaire 1846), 
misidentification]. Brimley's (1938) rec- 
ords of Ceresa uniformis Fairmaire ac- 
tually refer to Hadrophallus boreal is; 
these include one female in NCDA from 
Balsam [Jackson County], 14-18 Sep- 
tember. 

Stictocephala bisonia Kopp and Yonke 
1977 [as Ceresa bubalus (Fabricius 
1794), misidentification, in Brimley 
1938; as Stictocephala bubalus (Fabri- 
cius), misidentification, in Kopp and 
Yonke 1973b; Stictocephala bisonia 
Kopp and Yonke 1977 (Kopp and Yonke 
1977); as S. aha (Walker 1851) in An- 
drade 1997 (questionable synonymy, see 
"Notes," below)]. Counties: Alleghany, 
Ashe, Avery. Beaufort, Buncombe, Hay- 
wood, Madison, Mitchell, Montgomery, 
Stanly, Surry, Wake, Warren, Watauga. 
Seasonal distribution: 16 July-30 Sep- 
tember Host associations: Glycine max. 
Notes: This species is commonly known 
as the "buffalo treehopper" Recently, 
Andrade (1997) considered S. bisonia to 
be a junior synonym of 5". alta (Walker 
1851). We believe this synonymy is in- 
sufficiently justified in light of the need 
to examine the male genitalia to reliably 
identify many species of the tribe Cere- 



VOLUME 101, NUMBER 2 



255 



sini. The holotype of Ceresa alta Walker 
is of ambiguous identity, being a female 
specimen from an unspecified locality. 
On the other hand, the identity of the 
male holotype of S. bisonia Kopp and 
Yonke is definitive, and this type is ac- 
companied by 45 paratypes (males and 
females), all from Columbia, Missouri, 
USA. 

{Stictocephala brevicornis (Fitch 1856)]. 
Records of this species in North Carolina 
were not verified by the present authors. 
Brimley's (1938) record of Ceresa brev- 
icornis Fitch (Swannanoa, Buncombe 
County, May) probably referred to Stic- 
tocephala brevity his (Van Duzee); Kopp 
and Yonke's (1973b: map) record may be 
based on Brimley's publication. Hargrove 
(1986) also listed this treehopper from 
Coweeta, Macon County, on Robinia 
pseudoacacia. 

[Stictocephala brevis (Walker 1851)]. 
Notes: Brimley's (1938) record of this 
species in North Carolina (as Ceresa 
brevis Walker) has not been verified by 
the present authors. One specimen 
(Swannanoa, Buncombe County, 21 
May: NCDA) identified as Ceresa brevis 
Walker by Brimley is actually Stictoceph- 
ala brevitylus, however, Brimley's pub- 
lished record gave no locality or date. 

Stictocephala brevitylus (Van Duzee 1908) 
[as Ceresa brevitylus Van Duzee in Brim- 
ley 1938]. Counties: Alleghany, Anson, 
Avery, Brunswick, Buncombe, Cabarrus, 
Caswell, Catawba, Chatham, Cumber- 
land, Currituck, Dare, Durham, Forsyth, 
Granville, Harnett, Haywood, Henderson, 
Hertford, Jackson, McDowell, Mecklen- 
burg, Moore, New Hanover, Pasquotank, 
Randolph, Stanly, Swain, Wake, Wash- 
ington, Watauga, Wilkes, Yadkin, Yan- 
cey. Seasonal distribution: 5 April-2 
July. Host associations: Aster sp., Cea- 
nothus sp.. Chrysanthemum leucanthe- 
mum, Helianthus sp.*. Moras sp., Quer- 
cus falcata, Robinia pseudoacacia, Ru- 
bus argutus*, Sarracenia flava, Smilax 
sp.*, Solanum tuberosum, Vaccinium sp. 



(as Polyc odium sp.), Vitis sp. 'French hy- 
brid'. 

Stictocephala diceros (Say 1824) [as Cer- 
esa diceros (Say) in Brimley 1938]. 
Counties: Ashe, Buncombe, Cabarrus, 
Caldwell, Durham, Granville, Haywood, 
Iredell, McDowell, Mecklenburg, Wake, 
Watauga. Seasonal distribution: 8 June-6 
October. Host associations: Sambucus 
canadensis. 

[Stictocephala diminuta Van Duzee 1908]. 
North Carolina records of this species 
were not verified by the present authors. 
Brimley's (1938) record of 5'. diminuta in 
Raleigh [Wake County], July, probably 
represents S. brevitylus. 

Stictocephala lutea (Walker 1851). Coun- 
ties: Alleghany, Avery, Bladen, Bruns- 
wick, Buncombe, Burke, Carteret, Cas- 
well, Chatham, Cumberland, Duplin, 
Haywood, Hoke, Jackson, Johnston, 
Madison, Mecklenburg, Moore, New 
Hanover, Onslow, Pender, Sampson, 
Scotland, Stanly, Transylvania, Vance, 
Wake, Washington, Watauga. Seasonal 
distribution: 24 February- 16 September. 
Host associations: Quercus falcata (suck- 
ers). 

Stictocephala militaris (Gibson and Wells 
1917). NEW STATE RECORD. Fig. 6. 
Counties: Bladen, Craven. Hyde, John- 
ston, Mecklenburg, Pitt, Wake. Seasonal 
distribution: 19 June- 16 October. Host 
associations: Cercis canadensis, Prunus 
serotina. 

Stictocephala palmeri (Van Duzee 1908) 
[as Ceresa palmeri Van Duzee in Brim- 
ley 1938]. Counties: Avery-Caldwell- 
Watauga [as "Grandfather Mountain," 
which is on the border of these three 
counties (NCSU)], Buncombe, Graham, 
Macon, McDowell- Yancey [as "Buck 
Creek Gap," which is on the border of 
these counties (NCSU)], Stanly, Wake. 
Seasonal distribution: 21 July-29 Sep- 
tember. Host associations: no data for 
North Carolina. Notes: Elsewhere, Carya 
is the oviposition host (Funkhouser 



256 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



1917) — feeding occurs also on other 
woody hosts (Kopp and Yonke 1973b). 

Stictocephala stimulea (Van Duzee 1914). 
NEW STATE RECORD. Counties: Ca- 
barrus, Pender, Wake. Seasonal distribu- 
tion: 1—31 May. Host associations: Vitis 
prob. vulpina [ovipositing in canes]. 

[Stictocephala siibstriata (Walker 1851)]. 
Brimley's (1938) records of this species 
in North Carolina (Raleigh [Wake Coun- 
ty] and Southern Pines [Moore County]) 
were not verified by the present authors. 

Stictocephala taurina (Fitch 1856) [as Cer- 
esa taurina (Fitch) in Brimley 1938]. 
Counties: Ashe, Avery, Buncombe, For- 
syth, Madison, Mitchell, Surry, Wake, 
Watauga. Seasonal distribution: 19-20 
June-18 September. Host associations: 
Helianthus sp., Riibus sp., Sambuciis can- 
adensis, Smilax sp.* 

Stictocephala tauriniformis Caldwell 1949. 
NEW STATE RECORD. Counties: Hay- 
wood. Seasonal distribution: 14 Septem- 
ber. Host associations: no data for North 
Carolina. Notes: Elsewhere, Kopp and 
Yonke (1973b) reported Qiiercus and Til- 
ia as hosts. 

Stictolobus minutus (Funkhouser 1915) [as 
Stictolobus subulatus (Say 1830) in 
Brimley 1938 and Kopp and Yonke 
1973b]. Counties: Pender, Wake. Season- 
al distribution: early July. Host associa- 
tions: no data for North Carolina. Notes: 
Elsewhere, Kopp and Yonke (1973b) re- 
ported Taxodiwn distichiim (L.) L. Ri- 
chard as a host. 

Telamona ampelopsidis (Harris 1841). 
Counties: unknown, as "NC" on data la- 
bel (USNM). Seasonal distribution: un- 
known. Host associations: no data for 
North Carolina. Notes: Early reports of 
this species in North Carolina (Ball 1931; 
Brimley 1938) apparently refer to T. ma- 
culata Van Duzee, based on a specimen 
misidentified by C. S. Brimley as T. am- 
pelopsidis (NCDA). Kopp and Yonke 
(1974: map), who also reported the spe- 
cies in North Carolina, listed its host as 



Parthenocissus quinquefolia (L.) Plan- 
chon. 

Telamona collina (Walker 1851). NEW 
STATE RECORD. Counties: Wake. Sea- 
sonal distribution: 18-22 May. Host as- 
sociations: Platanus occidentalis. 

[Telamona concava Fitch 1851, misidenti- 
fication]. Records of Telamona concava 
from Linville Falls (Caldwell County, 
June 1920: NCDA) (Brimley 1942, Wray 
1967), actually refer to Heliria gibberata. 

Telamona decorata Ball 1903. NEW 
STATE RECORD. Counties: Ashe, Hen- 
derson, Watauga. Seasonal distribution: 
19-20 June- 13 September. Host associ- 
ations: Castanea dentata, Quercus alba, 
Q. rubra. 

Telamona dubiosa Van Duzee 1916. NEW 
STATE RECORD. Counties: Bladen. 
Seasonal distribution: 21 September-11 
October. Host associations: no data for 
North Carolina. Funkhouser (1917) listed 
Quercus alba as a host. 

Telamona extrema Ball 1903. NEW STATE 
RECORD. Counties: Carteret, Wake, 
Yadkin. Seasonal distribution: 13-29 
May. Host associations: no data for North 
Carolina. Notes: Hosts reported else- 
where include two species of Quercus 
(Kopp and Yonke 1974). 

Telamona maculata Van Duzee 1908. NEW 
STATE RECORD. Counties: Ashe, 
Wake. Seasonal distribution: 10 May- 18 
July. Host associations: no data for North 
Carolina. Notes: Hosts elsewhere include 
three species of Quercus (Kopp and Yon- 
ke 1974). 

Telamona monticola (Fabricius 1803). 
Counties: Ashe, Avery, Buncombe, 
Burke, Caldwell, Columbus, Hoke, John- 
ston, Macon, Robeson, Surry, Vance, 
Wake, Watauga. Seasonal distribution: 20 
April— 18 October. Host associations: 
Quercus falcata, Q. margarettiae, Q. ni- 
gra*, Q. rubra*, Q. stellata, Vitis rotun- 
difolia. 

Telamona reclivata Fitch 1851. Counties: 
Ashe, Buncombe, Macon, Mecklenburg, 



VOLUME 101, NUMBER 2 



257 



Wake. Seasonal distribution: 2 June-31 
August. Host associations: Quercus alba. 

[Telamoua salvini Distant 1879]. Records 
(Ball 1931, Brimley 1938) of T. salvini 
in North Carolina (Black Mountains) 
were not verified by the present authors, 
but the species is known to occur in 
Charleston County, South Carolina 
(NCSU, 1 specimen). 

[Telamona tiliae Ball 1925]. Records of T. 
tiliae in "North Carolina" (Ball 1931, 
Brimley 1938) seem to be based, at least 
in part, on a specimen (Blowing Rock 
[Watauga County], 13 Sept., NCDA) 
identified by C. S. Brimley as T. tiliae 
which is actually T. decorata. Presence 
of this species in North Carolina remains 
unconfirmed. 

Telamona unicolor Fitch 1851. Counties: 
Bertie, Bladen, Buncombe- Yancey [as 
"Black Mountains" which is on the Bun- 
combe-Yancey border], Columbus, 
Hoke, Macon, Mecklenburg, Wake. Sea- 
sonal distribution: 29 April-27 October. 
Host associations: Carya illinoinensis*, 
C. sp.*. 

Telamona westcotti Coding 1893. Counties: 
Henderson, Union. Seasonal distribution: 
25-30 June-5 September. Host associa- 
tions: no data for North Carolina. Notes: 
Hosts reported elsewhere include Quer- 
cus, Tilia, and Ulmus (Kopp and Yonke 
1974). 

Thelia bimaculata (Fabricius 1794). Coun- 
ties: Alexander, Ashe, Avery, Buncombe, 
Graham, Haywood, Jackson, Lincoln, 
Macon, McDowell, Mecklenburg, Swain, 
Wake, Watauga. Seasonal distribution: 10 
June-19 October. Host associations: Ro- 
binia pseudoacacia*. 

Tortistilus abnormus (Caldwell 1949). 
NEW STATE RECORD. Counties: Dur- 
ham, Stanly. Seasonal distribution: 13-21 
July. Host associations: no data for North 
Carolina. 

[Tortistilus inermis (Fabricius 1775)]. 
Brimley 's (1938) record of T. inermis (as 
Stictocephala inermis) in North Carolina 
was not verified by the present authors. 



but the species has also been recorded 
from two adjoining states, Virginia and 
Tennessee (Kopp and Yonke 1973b). 

Tortistilus lateralis (Funkhouser 1936). 
NEW STATE RECORD. Counties: Blad- 
en. Seasonal distribution: 8-22 Septem- 
ber. Host associations: no data for North 
Carolina. 

Tylopelta gibbera (Stal 1869) [as Tylopelta 
brevis Van Duzee 1908, in Brimley 1938; 
as Tylopelta americana (Coding 1893) in 
Kopp and Yonke 1973a]. Counties: Bun- 
combe, Haywood, New Hanover, Swain, 
Wake. Seasonal distribution: 16 April-27 
September. Host associations: Desmo- 
dium sp. 

Vanduzea arquata (Say 1830) [as V. arcu- 
ata (sic) in Brimley 1938]. Fig. 8. Coun- 
ties: Alamance, Alexander, Alleghany, 
Ashe, Avery, Bladen, Buncombe, Cald- 
well, Caswell, Catawba, Chatham, Cher- 
okee, Clay, Durham, Forsyth, Graham, 
Haywood, Jackson, Lincoln, Macon, 
Madison, Mecklenburg, Mitchell, 
Orange, Polk, Richmond, Rockingham, 
Sampson, Surry, Swain, Union, Wake, 
Watauga, Wilkes, Yancey. Seasonal dis- 
tribution: [? late March (NCSU)], 9 
May-27 October. Host associations: Ro- 
binia pseudoacacia*. 

Vanduzea triguttata (Burmeister 1836). 
Counties: Hoke, Johnston, Lee, Moore, 
New Hanover, Pender, Stanly, Wake. Sea- 
sonal distribution: 17 June-27 Septem- 
ber. Host associations: Lespedeza sp.* 

Xantholobus intermedins (Emmons 1854). 
Counties: Ashe, Buncombe- Yancey [as 
"Valley of Black Mountains," which is on 
the Buncombe- Yancey border (USNM)], 
Wake. Seasonal distribution: 4 May-7 
July. Host associations: Betula alleghan- 
iensis (as B. lutea).. Quercus phellos*. 

Xantholobus lateralis (Van Duzee 1908) [as 
Cyrtolobus lateralis Van Duzee in Brim- 
ley 1938]. Counties: Ashe, Burke, Cald- 
well, Lee, Mecklenburg, Watauga. Sea- 
sonal distribution: 3 May-6 July. Host 
associations: Betula sp., Quercus alba. 

Xantholobus muticus (Fabricius Mil) [as 



258 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Cyrtolobus muticus (Fabricius) in Brim- 
ley 1938]. Counties: Camden, Chatham, 
Duplin, Forsyth, Franklin, Gates, Greene, 
Hertford, Lee, Moore, Nash, Northamp- 
ton, Randolph, Rutherford, Wake, War- 
ren, Wayne, Wilson. Seasonal distribu- 
tion: 18 April- 14 July. Host associations: 
Quercus alba*, Q. laevis, Q. prinus, Q. 
stellata*. 
Xantholobus nitidus (Van Duzee 1908). 
Counties: Bladen, Forsyth, Johnston, 
Moore, Sampson, Wake, Wilkes. Season- 
al distribution: 17 May- 19 June. Host as- 
sociations: no data for North Carolina. 

Host Plant Index for 
North Carolina Treehoppers 

Plant scientific name, common name(s) 
(Family): associated treehopper species. 

Ambrosia artemisiifolia L., ragweed (Aster- 
aceae): Acutalis tartarea, Entylia cari- 
nata*, Publilia concava*. 

Ambrosia sp., ragweed (Asteraceae): Enty- 
lia carinata, Publilia concava*. 

Arachis hypogaea L., peanut, common pea- 
nut, goober, groundnut, grass nut, earth 
nut, monkey nut, pindar (Fabaceae): Spis- 
si stilus festinus*. 

Arundinaria sp., bamboo, cane (Poaceae): 
Acutalis tartarea. 

Aster ericoides L., heath aster (Asteraceae): 
Spissistilus festinus. 

Aster sp., aster, Michaelmas daisy, star wort, 
frost flower (Asteraceae): Entylia cari- 
nata, Stictocephala brevitylus. 

Betula alleghaniensis Britton, yellow birch, 
gray birch (Betulaceae): Xantholobus in- 
termedins. 

Betula sp., birch (Betulaceae): Carynota 
marmorata, Platycotis vittata*, Xantho- 
lobus lateralis. 

Bidens bipinnata L., Spanish needles (As- 
teraceae): Entylia carinata. 

Bidens coronata (L.) Britton, beggar ticks, 
beggar's ticks (Asteraceae): Acutalis tar- 
tarea, Entylia carinata. 

Bidens sp., beggar ticks, beggar's ticks, bur 
marigold, water marigold, pitchforks. 



Spanish needles, stick-tights, tickseed 
(Asteraceae): Entylia carinata. 

Carya illinoinensis (Wangenh.) K. Koch, 
pecan (Juglandaceae): Carynota mera, 
Microcentrus caryae*, M. perditus, Smi- 
lia fasciata, Telamona unicolor*. 

Carya sp., hickory (Juglandaceae): Cary- 
nota mera*, Enchenopa binotata com- 
plex, Microcentrus caryae, Telamona 
unicolor*. 

Castanea dentata (Marshall) Borkh., Amer- 
ican chestnut (Fagaceae): Atymna casta- 
neae*, Platycotis vittata*, Telamona de- 
corata. 

Castanea pumila (L.) P. Miller, chinquapin 
(Fagaceae): Atymna castaneae* . 

Ceanothus sp., redroot (Rhamnaceae): Stic- 
tocephala brevitylus. 

Cercis canadensis L., eastern redbud, red- 
bud, Judas tree (Fabaceae): Enchenopa 
binotata complex*, Stictocephala mili- 
taris. 

Chrysanthemum leucanthemum L., ox-eye 
daisy, white daisy, marguerite, white- 
weed (Asteraceae): Stictocephala brevi- 
tylus. 

Conyza canadensis (L.) Cronquist, horse- 
weed, hogweed, butter weed (Asteraceae): 
Entylia carinata, Micrutalis calva. 

Dahlia sp., dahlia (Asteraceae): Entylia 
carinata. 

Desmodium sp., beggar lice, beggar ticks, 
beggar's ticks, tick trefoil, tick clover 
(Fabaceae): Tylopelta gibbera. 

Erechtites hieraciifolia (L.) Raf. ex DC, 
fire weed (Ranuculaceae): Entylia cari- 
nata*. 

Erigeron annuus (L.) Persoon, daisy flea- 
bane, sweet scabious, white-top, fleabane 
(Asteraceae): Micrutalis calva. 

Erigeron sp., fleabane (Asteraceae): Entylia 
carinata. 

Eupatorium capillifolium (Lam.) Small, 
dog-fennel (Asteraceae): Acutalis tarta- 
rea, Entylia carinata. 

Eupatorium pilosum Walter, thoroughwort, 
boneset (Asteraceae): Entylia carinata. 

Eupatorium sp., thoroughwort, boneset 



VOLUME 101. NUMBER 2 



259 



(Asteraceae): Entylia carinata, Publilia 
concava* . 

Fagus grandifolia J. F. Ehrhart, beech, 
American beech (Fagaceae): Ophiderma 
definita. 

Fagus sp., beech (Fagaceae): Microcentrus 
caryae, Platycotis vittata. 

Gleditsia triacanthos L., honey locust, 
sweet locust, honeyshuck (Fabaceae): 
Micrutalis calva*. 

Glycine max (L.) Merrill, soybean, soja 
bean, soya bean (Fabaceae): Entylia car- 
inata, Micrutalis calva, Spissistilus fes- 
tinus*, Stictocephala alta. 

Helianthus annuus L., sunflower, common 
sunflower, mirasol (Asteraceae): Entylia 
carinata. 

Helianthus sp., sunflower (Asteraceae): 
Acutalis tartarea, Entylia carinata, Pub- 
lilia concava, Spissistilus festinus, Stic- 
tocephala brevity lus*, S. taurina. 

Helianthus tuberosus L., Jerusalem arti- 
choke, girasole (Asteraceae): Entylia car- 
inata*. 

Juglans nigra L., black walnut (Juglanda- 
ceae): Enchenopa binotata complex*. 

Lespedeza sp., bush clover (Fabaceae): 
Spissistilus festinus*, Vanduzea trigutta- 
ta*. 

Liriodendron tulipifera L., tulip tree, tulip 
poplar, whitewood, yellow poplar (Mag- 
noliaceae): Enchenopa binotata complex. 

Medicago sativa L., alfalfa, lucerne (Faba- 
ceae): Campylenchia latipes, Spissistilus 
festinus*. 

Morus sp., mulberry (Moraceae): Stictoce- 
phala brevitylus. 

Phaseolus vulgaris L., common bean, kid- 
ney bean, green bean, snap bean, haricot, 
French bean, frijol, runner bean, string 
bean, salad bean, wax bean (Fabaceae): 
Spissistilus festinus*. 

Physalis sp., ground cherry (Solanaceae): 
Micrutalis malleifera. 

Platanus occidentalis L., eastern sycamore, 
sycamore, button wood, buttonball, 
American plane-(Platanaceae): Telamona 
collina. 

Prunus serotina J. F. Ehrhart, black cherry. 



wild black cherry, rum cherry, (Rosa- 
ceae): Stictocephala militaris. 

Quercus alba L., white oak (Fagaceae): Ar- 
chasia auriculata*, A. belfragei*, Atym- 
na querci*, Cyrtolobus auroreus*, C. 
dixianus, C. fuscipennis* , Cyrtolobus gri- 
seus, C. maculifrontis* , C. pallidifrontis, 
C. tuberosus*, C. vau*, Glossonotus uni- 
vittatus, Ophidenna evelyna, O. flava*, 
O. flavicephala, Platycotis vittata*, Te- 
lamona decorata, T. reclivata, Xantho- 
lobus lateralis, X. muticus*. 

Quercus coccinea Muenchh., scarlet oak 
(Fagaceae): Cyrtolobus fuliginosus, 
Ophiderma flavicephala, Smilia fasciata. 

Quercus falcata Michaux, southern red oak, 
Spanish oak, Spanish red oak (Fagaceae): 
Stictocephala lutea, Telamona monticola, 
Cyrtolobus arcuatus*, C. dixianus, C. fe- 
nestratus*, C. fuliginosus*, C. inermis*, 
C. maculifrontis, Glossonotus acumina- 
tus*, Ophiderma definita, O. evelyna*, O. 
flavicephala* , O. pubescens* , O. sala- 
mandra, Platycotis vittata*, Smilia ca- 
melus*, S. fasciata*, Stictocephala brev- 
itylus, S. lutea, Telamona monticola. 

Quercus incana Bartram, bluejack oak, up- 
land willow oak, bluejack, turkey oak, 
high-ground willow oak, sand jack (Fa- 
gaceae): Platycotis vittata*. 

Quercus laevis Walter, turkey oak, Catesby 
oak (Fagaceae): Cyrtolobus fenestratus, 
C. ovatus*, Platycotis vittata, Smilia ca- 
melus*, Xantholobus muticus. 

Quercus laurifolia Michaux, laurel oak, 
Darlington oak, laurel-leaved oak (Faga- 
ceae): Ophiderma definita. 

Quercus margarettiae Ashe ex Small, 
scrubby post oak (Fagaceae): Cyrtolobus 
fenestratus, C. tuberosus, Telamona mon- 
ticola, Platycotis vittata. 

Quercus marilandica Muenchh., blackjack 
oak, blackjack, jack oak (Fagaceae): Cyr- 
tolobus fenestratus*, C. inermis, C. ova- 
tus*, C. tuberosus, Ophiderma evelyna*, 
O. pubescens. 

Quercus nigra L., water oak, possum oak 
(Fagaceae): Archasia auriculata, Cyrtolo- 
bus fenestratus* , C. fuscipennis, C. toga- 



260 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



tus*, C. tuberosus, Ophidertna definita*, 
O. evelyna, O. flavicephala* , O. salaman- 
dra, Platycotis vittata*, Smilia camel us*, 
S. fasciata*, Telamona monticola*. 

Qiierciis palustris Muenchh., pin oak, 
Spanish oak (Fagaceae): Cyrtolobus cla- 
ms*, C. dixianus, C. fenestratus, C. fu- 
liginosus, C. fimkhouseri, Entylia cari- 
nata, Ophiderma definita, O. evelyna, O. 
flavicephala, O. salamandra, Platycotis 
vittata, Smilia fasciata*. 

Quercus phellos L., willow oak (Fagaceae): 
Archasia belfragei, Cyrtolobus arcuatus, 
C. clams*, C. fenestratus*, C. fuligino- 
sus, C. togatus*, Ophiderma definita*, O. 
evelyna*, O. flavicephala* , O. salaman- 
dra*, Platycotis vittata, Smilia camelus, 
S. fasciata*, Xantholobus intermedius* . 

Quercus prinus L., chestnut oak, rock 
chestnut oak, basket oak (Fagaceae): Cyr- 
tolobus auroreus, C. tuberosus, C. vau, 
Xantholobus muticus. 

Quercus rubra L. (see also var. ambigua, 
gray oak or northern red oak, below), red 
oak (Fagaceae): Cyrtolobus fenestratus*, 
C. pulchellus, C. togatus, Glossonotus 
univittatus, Ophiderma flava, O. sala- 
mandra, Platycotis vittata*, Telamona 
decorata, T. monticola. 

Quercus rubra var. ambigua (Gray) Fern., 
gray oak, northern red oak (Fagaceae): 
Ophiderma flavicephala, Platycotis vit- 
tata. 

Quercus stellata Wangenh., post oak (Fa- 
gaceae): Atymna querci*, Cyrtolobus dix- 
ianus*, C fuliginosus, C. griseus, C. 
inermis, C. pallidifrontis*, C. togatus, C. 
tuberosus, C vau, Ophiderma flavicepha- 
la, O. pubescens, O. salamandra, Platy- 
cotis vittata*, Smilia camelus, S. fascia- 
ta*, Telamona monticola, Xantholobus 
muticus*. 

Quercus velutina Lam., black oak, yellow- 
bark oak, quercitron (Fagaceae): Archas- 
ia auriculata* , Cyrtolobus fenestratus, C. 
fuliginosus, Ophiderma salamandra* , 
Platycotis vittata, Smilia fasciata. 

Quercus virginiana P. Miller, live oak, 
southern live oak (Fagaceae): Cyrtolobus 



fenestratus, C. tuberosus, Platycotis vit- 
tata*. 

Robinia pseudoacacia L., black locust, 
false acacia, yellow locust (Fabaceae): 
Enchenopa binotata complex*, Micrutal- 
is calva, Stictocephala brevitylus, Thelia 
bimaculata* , Vanduzea arquata*. 

Rubus argutus Link, blackberry (Rosa- 
ceae): Stictocephala brevitylus*. 

Rubus sp., bramble (Rosaceae): Stictoceph- 
ala taurina. 

Salix nigra Marshall, black willow (Sali- 
caceae): Micmtalis calva. 

Sambucus canadensis L., elderberry, Amer- 
ican elderberry, sweet elderberry, (Capri- 
foliaceae): Stictocephala diceros, S. taur- 
ina. 

Sarracenia flava L., yellow pitcher plant, 
trumpets, watches, biscuit-flower, trum- 
petleaf, umbrella-trumpets, huntsman's 
horn (Sarraceniaceae): Spissistilus festin- 
us, Stictocephala brevitylus. 

Silphium sp., rosinweed (Asteraceae): En- 
tylia carinata. 

Smilax sp., greenbrier, catbrier (Liliaceae): 
Stictocephala brevitylus* , S. taurina*. 

Solanum tuberosum L., potato, Irish potato, 
white potato (Solanaceae): Entylia cari- 
nata, Stictocephala brevitylus. 

Solidago sp., goldenrod (Asteraceae): Acu- 
talis tartarea, Campylenchia latipes, En- 
tylia carinata, Micmtalis calva, Publilia 
concava*, Spissistilus festinus. 

Vaccinium sp., blueberry, huckleberry, 
cranberry, bilberry (Ericaceae): Sticto- 
cephala brevitylus. 

Verbesina alternifolia (L.) Britton ex Kear- 
ney, wingstem, yellow iron weed (Aster- 
aceae): Entylia carinata. 

Viburnum prunifolium L., black haw, sweet 
haw, sheepberry, nanny-berry, stagbush 
(Caprifoliaceae): Enchenopa binotata 
complex. 

Vitis rotundifolia Michaux, muscadine 
grape, scuppemong grape, bullace grape 
(Vitaceae): Atymna querci, Entylia cari- 
nata, Micmtalis calva, Telamona monti- 
cola. 

Vitis sp., grape (Vitaceae): Entylia carinata. 



VOLUME 101. NUMBER 2 



261 



Vitis sp. 'French hybrid,' grape 'French hy- 
brid' (Vitaceae): Micru talis calva, Stic- 
tocephala brevitylus. 

Vitis prob. viilpina L., frost grape, winter 
grape, chicken grape (Vitaceae): Sticto- 
cephala stimulea. 

Acknowledgments 

The authors are grateful to the following 
individuals: for collecting treehoppers listed 
in this work — J. R. Baker, C. H. Bartlett, R. 
L. Blinn, B. K. Cassel, P. J. Crawford, J. R. 
Cryan, B. A. Harrison, H. H. Neunzig, W. 
L. Nicholson, R. L. Regnery, D. L. Ste- 
phan, S. J. Toth, Jr., J. D. Trexler, M. S. 
Wallace, and B. M. Wiegmann; for prepar- 
ing many specimens for the North Carolina 
State University Insect Collection — M. P. 
Sobanski; for allowing us to examine spec- 
imens in their care — S. E. Halbert and F W. 
Mead (FSCA); K. R. Ahlstrom (NCDA); R. 
L. Blinn (NCSU); and T. J. Henry, S. H. 
McKamey, Systematic Entomology Labo- 
ratory, USDA, and R. C. Froeschner, 
Smithsonian Institution (USNM); for assis- 
tance with plant identifications and botani- 
cal nomenclature — J. W. Floyd and J. W. 
Hardin (Department of Botany, North Car- 
olina State University); and for helpful 
comments on manuscript — R. L. Blinn, J. 
R. Cryan, W. F Gimpel, Jr., S. H. Mc- 
Kamey, H. H. Neunzig, D. J. Voegtlin, B. 
M. Wiegmann, and D. W. Webb. Y. W 
Chan and S. J. Toth, Jr., assisted in the prep- 
aration of Fig. 11; photographs are by C. 
H. Dietrich, except Fig. 2, which is by R. 
L. Blinn. This research was supported by 
the North Carolina Agricultural Research 
Service and NSF grant DEB-9726282. 

Literature Cited 

Andrade, G. S. de. 1997. Stictocephala alta (Walker 
1851) sp. rev., comb, n., the coirect name for the 
"buffalo treehopper," with S. hisonia Kopp & 
Yonke, 1977 as a new synonym, and notes on 
Hadrophallus bubalus (Fabricius 1794) comb. n. 
(Homoptera: Membracidae). Transactions of the 
American Entomological Society 123: 289-295. 

Ball, E. D. 1931. A monographic revision of the tree- 



hoppers of the tribe Telamonini of North America. 
Entomologica Americana 12: 1-69. 

Brimley, C. S. 1938. The insects of North Carolina, 
being a checklist of the insects of North Carolina 
and their close relatives. Division of Entomology, 
North Carolina Department of Agriculture, Ra- 
leigh, North Carolina. 560 pp. 

Brimley, C. S. 1942. Supplement to insects of North 
Carolina. Division of Entomology, North Carolina 
Department of Agriculture, Raleigh, North Caro- 
lina. 39 pp. 

Deitz, L. L. 1975. Classification of the higher catego- 
ries of the New World treehoppers (Homoptera: 
Membracidae). North Carolina Agricultural Ex- 
periment Station Technical Bulletin 225: [i-iv], 1- 
177. 

Deitz, L. L. 1989. Bibliography of the Membracoidea 
(Homoptera: Aetalionidae, Biturritiidae, Membra- 
cidae, and Nicomiidae) 1981-1987. North Caro- 
lina Agricultural Research Service Technical Bul- 
letin 290: 1-31. 

Deitz, L. L. and C. H. Dietrich. 1993. Superfamily 
Membracoidea (Homoptera: Auchenorrhyncha). L 
Introduction and revised classification with new 
family-group taxa. Systematic Entomology 18: 
287-296. 

Deitz, L. L. and D. D. Kopp. 1987. Bibliography of 
the Membracoidea (Homoptera: Aetalionidae, Bi- 
turritiidae, Membracidae, and Nicomiidae) 1956- 
1980. North Carolina Agricultural Research Ser- 
vice Technical Bulletin 284: [i-ii], 1-39. 

Deitz, L. L., J. W. Van Duyn, J. R. Bradley, Jr., R. L. 
Rabb, W. M. Brooks, and R. E. Stinner 1976. A 
guide to the identification and biology of soybean 
arthropods in North Carolina. North Carolina Ag- 
ricultural Experiment Station Technical Bulletin 
238: [i-vi], 1-264. 

Eunkhouser, W. D. 1917. Biology of the Membracidae 
of the Cayuga Lake Basin. Cornell University Ag- 
ricultural Experiment Station Memoir 11: 177- 
445. 

Eunkhouser, W. D. 1927. Membracidae. General Cat- 
alogue of the Hemiptera. Eascicle 1 . Smith Col- 
lege, Northampton, Massachusetts. 581 pp. 

Coding, E W. 1893. Bibliographical and synonymical 
catalogue of the described Membracidae of North 
America. Bulletin of the Illinois State Laboratory 
of Natural History 3: 391-482. 

Hargrove, W. W. 1986. An annotated species list of 
insect herbivores commonly associated with black 
locust, Rohinia pseiutoacacia, in the southern Ap- 
palachians. Entomological News 97: 36-40. 

Kartesz. J. T 1994. A synonymized checklist of the 
vascular flora of the United States, Canada, and 
Greenland. Vol 2. Thesaurus. The biota of North 
America. 2nd edition. The University of North 
Carolina Press, Chapel Hill, North Carolina, vii -I- 
816 pp. 



262 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Kopp, D. D. and T. R. Yonke. 1973a. The treehoppers 
of Missouri: Part L Subfamilies Centrotinae, Ho- 
plophorioninae, and Membracinae (Homoptera: 
Membracidae). Journal of the Kansas Entomolog- 
ical Society 46: 42-64. 

Kopp, D. D. and T. R. Yonke. 1973b. The treehoppers 
of Missouri: Part 2. Subfamily Smiliinae; tribes 
Acutalini, Ceresini, and Polyglyptini (Homoptera: 
Membracidae). Journal of the Kansas Entomolog- 
ical Society 46: 233-276. 

Kopp, D. D. and T. R. Yonke. 1973c. The treehoppers 
of Missouri: Part 3. Subfamily Smiliinae; tribe 
Smiliini. Journal of the Kansas Entomological So- 
ciety 46: 375-421. 

Kopp, D. D. and T. R. Yonke. 1974. The treehoppers 
of Missouri: Part 4. Subfamily Smiliinae; tribe Te- 
lamonini (Homoptera: Membracidae). Journal of 
the Kansas Entomological Society 47: 80-130. 

Kopp, D. D. and T R. Yonke. 1977. Taxonomic status 
of the buffalo treehopper and the name Ceresa 
huhalus. Annals of the Entomological Society of 
America 70: 901-905. 

Kopp, D. D. and T. R. Yonke. 1979. A taxonomic re- 
view of the tribe Ceresini (Homoptera: Membra- 
cidae). Miscellaneous Publications of the Ento- 
mological Society of America 1 1(2): 1-97. 

Leonard, M. D. 1928. Membracidae, pp. 147-156. //; 
Leonard, M. D., ed., A list of insects of New York 
with a list of the spiders and certain other allied 
groups. Cornell University Agricultural Experi- 
ment Station Memoir 101, 1121 pp. 

Liberty Hyde Bailey Hortorium. 1976. Hortus Third. 
A concise dictionary of plants cultivated in the 
United States and Canada. Macmillian, New York, 
xiv -H 1290 pp. 

McGiffen, K. C. and H. H. Neunzig. 1985. A guide to 
the identification and biology of insects feeding 
on muscadine and bunch grapes in North Caroli- 
na. North Carolina Agricultural Research Service 
Bulletin 470: [i-iv], 1-93. 

McKamey, S. H. 1998. Taxonomic catalogue of the 
Membracoidea (exclusive of leafhoppers): second 
supplement to Fascicle I — Membracidae of the 
General Catalogue of the Hemiptera. Memoirs of 
the American Entomological Institute 60: 1-377. 

Mead. E W. 1986. Micrutalis treehoppers and pseudo- 
curly top in Florida (Homoptera: Membracidae). 



Florida Department of Agriculture & Consumer 
Services, Division of Plant Industry, Entomology 
Circular 283: [i-iv]. 

Metcalf, Z. P. 1915. A list of the Homoptera of North 
Carolina. Journal of the Elisha Mitchell Scientific 
Society 31: 35-60. 

Metcalf, Z. R and V. Wade. 1963. A Bibliography of 
the Membracoidea and Fossil Homoptera (Ho- 
moptera: Auchennorhyncha [sic]). North Carolina 
State [University], Raleigh, iv -I- 200 pp. 

Metcalf, Z. P. and V. Wade. 1965. General Catalogue 
of the Homoptera. A Supplement to Fascicle I — 
Membracidae of the General Catalogue of the He- 
miptera. Membracoidea. In Two Sections. North 
Carolina State University, Raleigh. 1552 pp. 

Osborn, H. 1940. The Membracidae of Ohio. Ohio Bi- 
ological Survey Bulletin 7: 51-101. 

Pratt, G. and T. K. Wood. 1992. A phylogenetic anal- 
ysis of the Enchenopa hinotata species complex 
(Homoptera: Membracidae) using nymphal char- 
acters. Systematic Entomolology 17: 351—357. 

Pratt, G. and T. K. Wood. 1993. Genitalic analysis of 
males and females in the Enchenopa binotata 
(Say) complex (Membracidae: Homoptera). Pro- 
ceedings of the Entomological Society of Wash- 
ington 95: 574-582. 

Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. 
Manual of the vascular flora of the Carolinas. Uni- 
versity of North Carolina Press, Chapel Hill, 
North Carolina. Ixi -t- 1183 pp. 

Remes-Lenicov, A. M. M. de. 1973. Nota sinonimica 
sobre Entylia carinata (Forster) (Homoptera: 
Membracidae). Revista de la Sociedad Entomo- 
logica Argentina 34: 89-94. 

Van Duzee, E. P. 1917. Catalogue of the Hemiptera of 
America north of Mexico excepting the Aphidi- 
dae, Coccidae and Aleurodidae [sic]. California 
Agricultural Experiment Station Entomological 
Technical Bulletin 2: [i-xiv], 1-902. 

Wray, D. L. 1950. Insects of North Carolina. Second 
supplement. Division of Entomology, North Car- 
olina Department of Agriculture, Raleigh, North 
Carolina. 59 pp. 

Wray, D. L. 1967. Insects of North Carolina. Third 
supplement. Division of Entomology, North Car- 
olina Department of Agriculture, Raleigh, North 
Carolina. 181 pp. 



PROC. ENTOMOL. SOC. WASH. 
101(2). 1999. pp. 263-269 

A TURN OF THE CENTURY CONUNDRUM— REEXAMINATION OF 
AEOLOTHYNNUS ASHMEAD (HYMENOPTERA: TIPHIIDAE: THYNNINAE) 

Lynn S. Kimsey 

Bohart Museum of Entomology, Department of Entomology, University of California, 
Davis, CA 95616, U.S.A. (e-mail: lskimsey@ucdavis.edu) 



Abstract. — The identity of the Australian thynnine genus Aeolothynnus Ashmead has been 
the source of nomenclatural confusion since the early 1900's. Its identity is reevaluated 
relative to other related genera, species placements are reconsidered, and one new species, 
Aeolothynnus caliventer, from South Australia, is described. Asthenothynnus is discovered 
to be a new junior synonym of Aeolothynnus, and Turner's concept of Aeolothynnus is 
in reality synonymous with Thynnoturneria Rohwer New combinations of species in 
Aeolothynnus include: Thynnus beatrix Turner, Asthenothynnus deductor Turner, Thynnus 
generosus Turner, Asthenothynnus kurandensis Turner, Thynnus lactarius Turner, Asthen- 
othynnus lilliputianus Turner, Asthenothynnus mahtimus Turner, Asthenothynnus minutis- 
simus Turner, Asthenothynnus perkinsi Turner, Asthenothynnus pleuralis Turner, Thynnus 
pulcherrimus Turner, and Asthenothynnus vicarius Turner. 

Key Words: Tiphiidae, Hymenoptera, Thynninae, Aeolothynnus, Asthenothynnus, Aus- 
tralia 



Many genera of Thynninae were origi- 
nally based on one or a small number of 
species, and the original characterizations 
were obscure at best. One of the finest ex- 
amples of the confusion resulting from 
these inadequate descriptions can be seen in 
the taxonomic literature near the turn of the 
century, published by taxonomists at the 
British Museum and at the U.S. National 
Museum. This confusion was largely the re- 
sult of miscommunications, misinterpreta- 
tions and a bit of transatlantic competition. 
Two generic names, Aeolothynnus Ash- 
mead and Asthenothynnus Turner, were pro- 
posed for the same group of species. The 
valid generic name for these species is Aeo- 
lothynnus and Turner's mistaken concept of 
the genus Aeolothynnus is a very different 
entity later renamed Thynnoturneria Roh- 
wer, which in turn has been confused with 
Iswaroides Ashmead. 



Ashmead originally based Aeolothynnus 
on the new species multiguttatus. He de- 
scribed the species and genus simultaneous- 
ly in one of his notorious keys (1903). As 
a result of his habit of describing new gen- 
era and species in very brief keys, most 
subsequent authors confused the identity of 
Aeolothynnus. Although Ashmead did not 
clearly indicate the species as new, the ge- 
nus was monotypic, therefore as stated in 
the International Zoological Code (1985), 
Article 12, the generic description also ap- 
plied to the species. As a result. Turner's 
1908 statement that 'Ashmead gives Aeo- 
lothynnus multiguttatus Ashm., as the type 
of his genus but, as he has not given any 
description of that species, his name cannot 
stand."" is incorrect. To further confuse the 
situation, Rohwer (1910a) subsequently 
recognized Ashmead's designation of mul- 
tiguttatus and gave a new name to Turner's 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



concept of Aeolothynnus as Turnerella, 
stating that: 

''The characters given by Ashmead in 
his table of the genera of Thynnidae, 
are sufficient to satisfy the technical re- 
quirements so this species [Aeolothyn- 
nus nndtiguttatus Ashmead (nee Turn- 
er)] should date from that time and be 
accredited to Ashmead.'' 
''Turner considering that Aelothynnus 
(sic!) multiguttatus Ashm, was unde- 
scribed named Thynnus cerceroides Sm. 
as the type of Aelothynnus. Aelothyn- 
nus multiguttatus Sm. and Thynnus 
cerceroides are not congeneric, which 
leaves Aelothynnus Turn, without a 
name. For this genus the name Tur- 
nerella may be used."' 

Unfortunately the name Turnerella ran into 
problems of homonymy, and according to 
Turner (1911): 

"My identification of Ashmead' s genus, 
of which the type was undescribed, was 
incorrect, as has been pointed out by 
Mr. Rohwer, who renamed the genus 
Turnerella. That name, however, was 
used by Professor Cockerell for a ge- 
nus of bees; his paper was published 
in London on the same day as Mr. 
Rohwer' s paper was published in 
America, and I believe the name 
should be retained for the bee. I there- 
fore have to propose a new name for 
the genus." 

Turner never made a new generic descrip- 
tion for his concept of Aeolothynnus. The 
uncertainty of the situation caused Given 
(1959) to lament: 

"The genus Aeolothynnus was erected 
by Ashmead in 1903 with the genotype 
A. multiguttatus. The genus was then 
very poorly defined and has been fre- 
quently misinterpreted by subsequent 
workers." 

"Rohwer (1910a) published the first 
description of the genotype, Ashmead 



(1903) having given mere key distinc- 
tions. Rohwer (1910a) stated that T 
cerceroides and A. multiguttatus were 
not congeneric and therefore the genus 
Aeolothynnus Turner was left without 
a name as that name was valid for the 
genotype multiguttatus of Ashmead. 
Rohwer (1910a) proposed the name 
Turnerella for Turner's genus. How- 
ever, this generic title was preoccu- 
pied, and both Turner (1911) and Roh- 
wer (1910b) appreciated this at about 
the same time. Turner (191 1) then pro- 
posed the name Eurohweria for his ge- 
nus, but he was forestalled by Rohwer 
(1910b) who proposed the name Thyn- 
noturneria." 

When Rohwer (1910b) renamed A^o/o/M-n- 
nus, as treated by Turner, he established 
Turner's concept of the group as a valid ge- 
nus, particularly since none of the species 
placed by Turner (1910a) in Aeolothynnus 
under cerceroides were congeneric with 
multiguttatus. Thus the name Thynnotur- 
neria Rohwer applied to the cerceroides 
group of species. Turner (1912) was not en- 
tirely pleased by this situation: 

"/ am by no means sure that the name 
Aeolothynnus should not be used for 
this genus. Ashmead in describing the 
genus Aeolothynnus took an undescribed 
species for the type. In my work on the 
Thynnidae 1 accepted Ashmead' s ge- 
nus, but treated the species as a nomen 
nudum. Mr. Rohwer, on the other 
hand, holds that the description of the 
genus covers the species also; but I 
cannot agree with this opinion, as Ash- 
mead evidently did not intend the de- 
scription for a specific one, and a de- 
scription to be recognized should be at 
least intended by the author for a de- 
scription of a species." 
"Unfortunately, A. cerceroides, Sm., 
selected by me as the type of the genus, 
does not appear to belong to the same 
genus as Ashmead' s type. Yet if Ash- 
mead's specific name is treated as a 



VOLUME 101. NUMBER 2 



265 



nomen nudum, A. cerceroides must be 
treated as the type of the genus. It is 
bad enough to have to recognized the 
very insufficient descriptions of some 
authors as valid, but if we are also to 
accept what were never intended for 
descriptions things would be still 
worse. For the present, pending some 
decision on the subject, I am using 
Rohwer's name, but do not consider 
that it can stand. The whole confusion 
is due to a want of editing in Ash- 
mead's paper, as no editor should pub- 
lish a description of a genus with an 
undescribed species taken for the 
type." 

Turner's argument that Ashmead did not in- 
tend to describe multiguttatus as new at the 
same time as he described the genus was 
not accepted by other systematists. 

Simultaneously, Turner (1910a) also 
named a new genus Asthenothynnus, based 
on Thynnus pulchellus Klug. Upon exami- 
nation, Thynnus pulchellus turns out to be 
congeneric with multiguttatus. Therefore 
Asthenothynnus also becomes synonymous 
with Aeolothynnus. In light of this generic 
confusion all of the species placed various- 
ly in Aeolothynnus, Thynnoturneria and As- 
thenothynnus need to be reevaluated for 
their correct placement. Some of this re- 
placement of species has been done below, 
showing new combinations as indicated. 
Types that have been seen and the generic 
placement confirmed are indicated by an as- 
terisk (*). However, those without either in- 
dication are placed in the appropriate genus 
based on whether Turner himself placed 
them in his concept of Aeolothynnus or one 
of the subsequent generic names for that en- 
tity, or placed them in Asthenothynnus, 
which is the equivalent of Ashmead's Aeo- 
lothynnus. 

To further clarify Aeolothynnus Ashmead 
the genus is rediagnosed and discussed be- 
low, species placements are reassigned, and 
a new species, caliventer, is described. 



which exhibits some very unusual thoracic 
modifications in the male. 

Materials and Methods 

Specimens were studied in situ or were 
borrowed from the following institutions: 
the Natural History Museum, London, S. 
Lewis; Hope Museum, Oxford University, 
C. O'Toole; and the Australian National In- 
sect Collection, CSIRO, Canberra, ACT, I. 
Naumann and J. Cardale. 

Aeolothynnus Ashmead 
(Figs. 1-10) 

Aeolothynnus Ashmead 1903: 101. Type 
species: Aeolothynnus multiguttatus Ash- 
mead 1903: 101. Monobasic. 

Asthenothynnus Turner 1910a: 34. Type 
species: Thynnus pulchellus Klug 1842: 
20. Original designation. New synony- 
my. 

Aelothynnus Rohwer 1910a. Invalid emen- 
dation of Aeolothynnus Ashmead 1903. 

Male. — Body length 3-10 mm. Head: 
face flattened in profile, interantennal area 
nearly planar with clypeus; antennal sockets 
with dorsal antennal lobe highly reduced 
without carinae or transverse welt; suban- 
tennal area highly polished and impunctate; 
apical 7 to 9 flagellomeres somewhat ar- 
cuate or bulging along inner margin, with 
two small linear tyloids; clypeus nearly flat 
or slightly convex in profile, apical margin 
narrowly truncate; labrum small, medially 
emarginate with narrow neck-like base; 
tongue short unmodified, prementum with 
apical brush of long setae reaching back to 
occipital foramen; maxillary stipes with 
short stipal fringe originating on inner mar- 
gin and extending transversely across stipes 
to outer margin; mandibles slender with 
small subapical tooth on inner margin; oral 
fossa extending to inner margin (Fig. 8) of 
mandible, widely separated from occipital 
carina; venter of head with large flattened 
and polished area, margined by carinae and 
surrounding the oral fossa. Thorax: prono- 
tum with transverse anterior carina well-de- 



266 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




3. caliventer 



4. caliventer 



5. caliventer 



6. 7. 

wesPtvoodi multigiittatus 



8. multiguttatus 



1 0. caliventer 



Figs. 1-10. 1-5, 9-10, Aeolothynniis caliventer. 6, A. westwoodi. 7-8, A. multiguttatus. 1, Ventral view of 
male thorax. 2, Lateral view of body. 3, Ventral view of genital capsule. 4, Lateral view of genital capsule. 5- 
7, Apex of hypopygium. 8, Ventral view of male head. 9, Posterior view of apex of female abdomen. 10. Male 
apical gastral tergum. 



veloped; mesepi sternum and mesopleuron 
separated by shallow scrobal groove; pro- 
podeum without discrete dorsal surface, 
gently sloping posteriorly; fore femur often 
indented basoventrally; forecoxae globular 
in most species; hindcoxa with short dorsal 
carina. Abdomen: basal segment slightly 
convex ventrally, foreshortened, dorsally 
without discrete dorsal surface; apical ter- 
gum hood-like, with narrow apical lip (Fig. 
10); hypopygium ventrally longitudinally 
grooved or trough-like, apically variable, 
tridentate, ligulate or unidentate (Figs. 5-7); 
abdominal segments smoothly tapering one 
to the next (Fig. 2); terga II-IV with fine 
sublateral transverse sulcus; spiracles not 



apparent; terga with W-shaped transverse 
sulcus or line, marked by a row of setae 
near posterior margin. Genital capsule 
(Figs. 3, 4): penis valves large, dilated api- 
cally and often curved; volsella large flat- 
tened and tapering apically, forming floor 
of capsule; digitus not apparent; parameres 
slender and subtriangular; gonocoxa pro- 
duced into elongate, often apically notched 
dorsal lobe (Fig. 3); aedeagus short, origi- 
nating on this lobe (Fig. 3), with short api- 
cal loop. Color: body black with yellow, 
white and red markings; vertex with oblong 
reddish brown spot extending diagonally 
from dorsal eye margin to behind hindo- 
celli; abdomen in most species with odd 



VOLUME 101, NUMBER 2 



267 



comma-shaped pale markings on most seg- 
ments. 

Female. — Body length 2-5 mm; Head: 
broader than long or elongate and usually 
appearing pinched across at eyes; clypeus 
narrow and linear, shorter than interantennal 
distance; maxillary palpus with two articles, 
labial palpus with three; mandible edentate 
and sickle-shaped. Thorax: pronotum sub- 
quadrate; forecoxa unmodified or narrowed 
and separated by deep rectangular slot 
{miiltiguttatus); scutum and metanotum 
broadly visible dorsally. Abdomen: tergum 
I and III-V with broadly W-shaped trans- 
verse sulcus; tergum II with three transverse 
carinae or ridges; tergum V with thin sha- 
greened or roughened laterotergite marked 
by an arcuate bulge and sulcus; tergum VI 
with broadly or narrowly ovoid plate delim- 
ited at least laterally by carina, subtended 
by long tuft of setae (Fig. 9); sternum VI 
narrowly U-shaped. Color: pale brown. 

Distribution. — This genus occurs 
throughout southern Australia in New 
South Wales, South Australia, Western Aus- 
tralia, Tasmania, Victoria, and apparently 
the Northern Territory, although this record 
needs to be confirmed. 

Included species. — Thirty species are 
placed in Aeolothynnus including: beatrix 
(Turner) 1908* (Thymius), new combina- 
tion; caliventer Kimsey, new species; de- 
coratus (Smith) 1879 (Thynnus)\ deductor 
(Turner) 1910b (Asthenothynniis), new 
combination; exiguus (Turner) 1910c 
(Thynuus); generosus (Turner) 1908* 
(Thynnus), new combination; incensus 
(Smith) 1868 (Thynnus); innociius (Turner) 
1908 (Thynnus); kurandensis (Turner) 
1910d* (Asthenothynnus), new combina- 
tion; lactarius (Turner) 1910d* (Thynnus), 
new combination; leucostictus (Turner) 
1908 (Thynnus); lilliputianus (Turner) 
1915a* (Asthenothynnus), new combina- 
tion; maritimus (Turner) 1915b* (Astheno- 
thynnus), new combination; minutissimus 
(Turner) 1910c (Asthenothynnus), new 
combination; minutus (Smith) 1859 (Thyn- 
nus); multiguttatus Ashmead 1903*; pene- 



tratus (Smith) 1879 (Thynnus); perkinsi 
(Turner) 1910d* (Asthenothynnus), new 
combination; planiventris (Turner) 1908 
(Thynnus); pleuralis (Turner) 1915a (As- 
thenothynnus), new combination; pulchel- 
his (Klug) 1842 (Thynnus) ( = Thynnus mul- 
tipictus Smith 1879); pulcherrimus (Turner) 
1908* (Thynnus), new combination; pyg- 
maeus (Turner) 1908 (Thynnus); quadricar- 
inatus (de Saussure) 1867 (Thynnus); rub- 
romaculatus (Turner) 1908 (Thynnus); ten- 
uis (Turner) 1908 (Thynnus); vicarius 
(Turner) 1915a (Asthenothynnus), New 
combination; westwoodi (Guerin de Mene- 
ville) 1842 (Agriomyia) ( = Thynnus intri- 
catus Smith* 1859); longiceps (Smith) 
1859 (Thynnus); nanus (Smith) 1879 
(Thynnus). 

Discussion. — Members of the genus 
Aeolothynnus are small-bodied and locally 
abundant Australian wasps. The vast ma- 
jority of species average 1 cm in length or 
less. Thousands of individuals, both males 
and females, may be found on a single 
flowering Eucalyptus tree. Members of the 
genus occur in most habitats throughout at 
least the southern part of Australia. They 
are for the most part unremarkably modi- 
fied. However, a new species, collected in 
South Australia has a bizarrely modified 
male. The male modifications in this spe- 
cies are unusual for the entire subfamily, so 
it is described as new below. Hosts are ap- 
parently unknown for Aeolothynnus. How- 
ever, given the parasitic habits of the rest of 
the subfamily, the hosts are undoubtedly 
small, locally abundant, species of larval 
Scarabaeidae. 

A number of traits are diagnostic for 
members of this genus. The most distinctive 
feature of the males is the longitudinally 
grooved or impressed apical abdominal 
sternum (hypopygium). This characteristic 
coupled with their smoothly tapering ab- 
domen, and flattened face will distinguish 
Aeolothynnus males from closely related 
genera. Aeolothynnus belongs to a group of 
genera characterized by the presence of a 
transverse carina, ridge or welt, across the 



268 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



apical edge of the apical abdominal tergum 
(epipygium), the female apical abdominal 
tergum is smoothly convex, narrowed, with 
an oval medial plate margined laterally by 
a carina, which is in turn subtended later- 
ally by a long brush of setae. This group 
consists of Epactiothynnus Turner, Tmeso- 
thynnus Turner, Thynnoturneria Rohwer, Is- 
waroides Ashmead, Gymnothynnus Turner, 
Acanthothynnus Turner, Doratithynniis 
Turner and Aspidothynnus Turner. Females 
are problematic in this group of genera and 
too few are associated with males to distin- 
guish between species level and generic 
characteristics. 

Aeolothynnus caliventer Kimsey, 
new species 

(Figs. 1-5, 9, 10) 

Male. — Body length 8-9 mm; Head: 
face nearly flat from frons to upper clypeus; 
frons and vertex densely punctate, punc- 
tures small and contiguous; prementum 
strongly convex medially, with long medial 
fringe of setae; flagellum I 1.5X as long as 
broad; flagellum II 2.5 X as long as broad; 
flagellum III three times as long as broad. 
Thorax (Figs. 1, 2): Pronotum extended 
ventrally; propleura convex; forecoxa ven- 
trally flat and expanded sharp-edged later- 
ally; forefemur flat, broadly expanded and 
rounded ventrally; mesepistemum strongly 
produced and flange-like, giving the thorac- 
ic venter a strongly cuplike appearance. Ab- 
domen: epipygium (Fig. 10); hypopygial 
apex subtruncate with rounded lateral cor- 
ners and short acute medial spine, ventral 
groove teardrop-shaped (Fig. 5). Genital 
capsule (Figs. 3, 4). Color: body black with 
pale whitish markings on apical margin of 
clypeus, a small spot on each antennal lobe, 
pronotum with small spot adjacent to tegula 
and on either end of transverse anterior ca- 
rina; small spot at posterior angle of me- 
sopleuron above midcoxa outlining meso- 
pleural lamellae, and small comma-shaped 
lateral spot on gastral terga I-VI; vertex 
with small oval red spot between hindocelli 
and nearest eye margin; fore- and midlegs 



with femoral apex and tibia entirely or part- 
ly red; hindfemur medially red; hindtibia 
somewhat reddish medially; wing mem- 
brane untinted, veins dark brown. Pubes- 
cence: long erect and silvery, except dense 
and golden along mesepisternal edge. 

Female. — Body length 5.5 mm; Head: 
head broader than long; mandible sickle- 
shaped and edentate. Thorax: pronotal disk 
with anterolateral comers acute, posterior 
margin with two submedial warts; propleu- 
ra strongly convex ventrally; scutum broad- 
ly visible; scutellum flattened, length sub- 
equal to breadth; propodeum strongly con- 
vex and rounded laterally and posteriorly. 
Abdomen: tergum I with W-shaped sulcus; 
II with three transverse ridges; III-IV with 
large submedial U-shaped sulcus; pygidium 
narrowly longitudinally ovoid with lateral 
carina, with row of setae laterad of carina 
(Fig. 9); sternum VI apex narrowly U- 
shaped. Color: yellowish brown. 

Material examined. — Holotype S: Aus- 
tralia: SA, 79 km nnw Renmark, 33°3rS 
140°24'E, 9 Aug.-7 Sept. 1995, K. R. Pul- 
len, Casuarina woodland, malaise trap. Ho- 
lotype deposited in the Australian National 
Insect Collection (ANIC), Canberra, ACT. 
Paratypes: two J, one 9, same data as ho- 
lotype except also collected in flight inter- 
cept trap (deposited in ANIC and Bohart 
Museum of Entomology, University of Cal- 
ifornia, Davis). These specimens derived 
from the Calperum Station/Bookmark Bio- 
sphere Reserve Invertebrate Survey. 

Etymology. — The name refers to the pe- 
culiar modifications of the male thorax; ca- 
lix = cup, venter = belly, Latin, noun. 

Discussion. — The unusually modified 
male thorax and forefemur will serve to dis- 
tinguish this species from other Aeolothyn- 
nus or related genera. The female is less 
remarkable but can be distinguished by 
shape of the pronotum with a small acute 
tooth on the anterolateral corner and pos- 
terior submedial swellings. In addition, the 
scutellum is relatively flat and the propleura 
are strongly bulging ventrally. 



VOLUME 101, NUMBER 2 



269 



Acknowledgments 

Many thanks to Richard M. Bohart for 
reviewing the manuscript, and to the vari- 
ous collections and curators who made the 
study possible. 

Literature Cited 

Ashmead, W. H. 1903. Classification of the fossorial, 
predaceous and parasitic wasps, of the superfam- 
ily Vespoidea. Canadian Entomologist 35: 95- 
107. 

Given, B. B. 1959. Notes on Australian Thynninae. V. 
Aeolothynnus miiltiguttatiis Ashmead and Iswa- 
roides koebelei Ashmead. Proceedings of the Lin- 
naean Society of New South Wales 83: 400-402. 

Guerin de Meneville, F. E. 1842. Materiaux sur les 
Thynnides. Magasin de Zoologie 4: 1-15. 

International Code of Zoological Nomenclature. 1985. 
Third edition. University of California Press, 
Berkeley, xx + 338 pp. 

Klug, J. C. E 1842. Ueber die Insectenfamilie Heter- 
ogyna Latr. und die Gattung Thynnus E insbeson- 
dere. Physikalische und Mathematische Abhan- 
dlungen der Koniglichen Akademie der Wissen- 
schaften zu Berlin, pp. 1-44. 

Rohwer, S. A. 1910a. Turner's genera of Thynnidae 
with notes on Ashmeadian genera. Entomological 
News 21: 345-351. 

. 1910b. A preoccupied generic name (Tliyn- 

uoturneha n. n.). Entomological News 21: 474. 

de Saussure, L. E 1867. Hymenoptera. //; Reise der 
Osterreichischen Eregatte Novara uni die Erde in 
den Jahren, etc. Zoologischer Theil, Zweiter 
Band: Hymenoptera. Vol. 2, 156 pp. 

Smith, E 1859. Catalogue of the Hymenopterous in- 



sects in the collection of the British Museum 7: 
10-69. 

. 1868 (May). Descriptions of aculeate Hyme- 
noptera from Australia. Transactions of the Ento- 
mological Society of London 1868: 231-237. 

. 1879. Descriptions of new species of Hyme- 
noptera in the British Museum, London. 240 pp. 

Turner, R. E. 1908. A revision of the Thynnidae of 
Australia. Pt. II. Proceedings of the Linnaean So- 
ciety of New South Wales, Sydney 33: 70-208 
(June), 209-256 (August). 

. 1910a. Hymenoptera, Family Thynnidae. In 
Wytsman, P. ed.. Genera Insectorum, 105. Brux- 
elles, 62 pp. 

. 1910b. Additions to our knowledge of the fos- 



sorial wasps of Australia. Proceedings of the Zoo- 
logical Society of London 1910: 259-306. 

. 1910c. New species of Thynnidae from the 

Australian and Austro-Malayan Regions. Annales 
Historico-Naturales Musei Nationalis Hungarici 8: 
107-124. 

. 1910d. New fossorial Hymenoptera from Aus- 
tralia. Transactions of the Entomological Society 
of London 1910: 407-429. 

. 1911. Notes on fossorial Hymenoptera. V. 

Further notes on the Thynnidae and Scoliidae. An- 
nals and Magazine of Natural History, London 
(8)8: 602-624. 

. 1912. Notes on fossorial Hymenoptera, IX. 

On some new species from the Australian and 
AustroMalayan Regions. Annals and Magazine of 
Natural History, London (8)10: 48-63. 

. 1915a. Descriptions of new fossorial wasps 

from Australia. Proceedings of the Zoological So- 
ciety of London 1915: 41-69. 

. 1915b. Notes on fossorial Hymenoptera. XVI. 

On the Thynnidae, Scoliidae and Crabronidae of 
Tasmania. Annals and Magazine of Natural His- 
tory (8)15: 537-559. 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999. pp. 270-273 

NEW NORTH AMERICAN SPECIES AND RECORDS 

IN THE GENUS XENOLIMOSINA ROHACEK 

(DIPTERA: SPHAEROCERIDAE: LIMOSININAE) 

S. A. Marshall 

Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada 
NIG 2W1 



Abstract. — Xenolimosina glabrigena, new species, is described from northern Florida, 
the female of Xenolimosina phoba Marshall is described, and new distributional records 
are given for X. phoba and X. sicula Marshall. 

Key Words: Sphaeroceridae, Diptera, Florida insects 



Xenolimosina Rohacek is one of the most 
rarely collected genera of Sphaeroceridae, 
probably because most species seem to be 
active only during the colder months of the 
year. Xenolimosina previously included 
three species: X. setaria Villeneuve from 
Europe, X. sicula Marshall from Ontario, 
Quebec, Arkansas and California, and X. 
phoba Marshall from Quebec and Mary- 
land. The latter species was previously 
known only from males, and a description 
of the female is given below along with 
new distributional records for both de- 
scribed North American species. A third 
Nearctic species, Xenolimosina glabrigena 
new species, is described from northern 
Florida. Xenolimosina glabrigena is recog- 
nisable as a Xenolimosina by the long ex- 
serted hind tibial bristle, small eyes, tele- 
scoping female abdomen, lack of a mid 
ventral bristle on the mid tibia, two spinose 
lobes on the male fifth sternite (Fig. 2), and 
the short surstylus with a long-setose pos- 
terolateral surface (Fig. 3). Xenolimosina 
glabrigena will key out to X. sicula in Mar- 
shall (1985), but it is distinctly different 
from X. sicula in details of the male and 
female genitalia. Most notably, the disti- 



phallus of X. glabrigena is simple, broad 
and tubular, much like X. phoba (Marshall 
1985, fig. 23), but in marked contrast to the 
broad, spinose and highly modified disti- 
phallus of X. sicula (Marshall 1985, fig. 22) 
and X. setaria (Rohacek 1983, fig. 322). 
Some of the characteristics of X. glabri- 
gena, such as the dense pile of the male 
fore tibia and basitarsus, the shape of the 
postgonite (Fig. 1), and the sclerotization of 
the terminal sclerites of the female abdo- 
men, are distinctive autapomorphies not 
found in the other species. 

Rohacek (1982) suggested that Xenoli- 
mosina belongs to the Minilimosina genus- 
group, and that it is probably the sister ge- 
nus to Minilimosina. The distiphallus in 
Minilitnosina species is generally simple 
and unadorned, suggesting that the charac- 
teristically adorned distiphallus of X. sicula 
and the European species X. setaria are 
synapomorphic, and these two species form 
a monophyletic group. The other two spe- 
cies in the genus, X. glabrigena and X pho- 
ba, both have an elongate tubular distiphal- 
lus (Fig. 1) and a greatly enlarged costagial 
bristle, and together probably form the sis- 
ter group to X. setaria plus X. sicula. 



VOLUME 101, NUMBER 2 



271 




Figs. 1-10. 1-7. Xenolimosina i^Uihrigena. 1, Aedeagus and associated structures. 2, Male sternites 5-7. 3, 
Male terminalia, left lateral. 4, Male terminalia, posterior 5, Female terminalia, dorsal. 6, Female terminalia, 
ventral. 7. Spermathecae. 8-10. X. phoba. 8. Spermathecae. 9. Female terminalia. dorsal. 10. Female terminalia, 
ventral. 



272 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Xenolimosina glabrigena Marshall, 
new species 

(Figs. 1-7) 

Description. — Body length ca. 2.0 mm, 
dark brown to black with a heavy pruinos- 
ity; lower frons and gena reddish; legs 
brown. Interfrontal plate 1.5X as high as 
width at middle, bordered by 4 pairs of 
equal interfrontal bristles, lower two pairs 
weakly cruciate. Postocellar bristles absent. 
Eye small, height ca. 1.3X genal height, 
gena shining except for lower margin and 
a narrow vertical strip on posterior third. 
Palpus strongly swollen at middle, tapered 
at apex. Scutum with 5-6 rows of acrosti- 
chal bristles between anterior dorsocentral 
bristles; dorsocentral bristles in 2 pairs, an- 
terior pair 0.6 X as long as posterior pair, 
posterior pair equal to scutellar length. 
Prescutellar acrostichal bristles in a single 
pair twice as long as acrostichal setulae. 
Foreleg of male with dense yellow pile ven- 
trally on distal half of tibia and on tarso- 
mere one; foreleg of female unmodified. 
Mid tibia with long proximal anterodorsal 
distal anterodorsal and distal dorsal bristles. 
Katepistemum pruinose, with a short pos- 
terodorsal bristle and a minute anterodorsal 
bristle. Wing length 2.5 X width; second 
costal sector 1.0-1.4X third; costa extend- 
ing about 2 vein-widths beyond apex of 
R4+V Costagial bristle very long, longer 
than alula; alula narrow. 

Male abdomen: Sternite 5 (Fig. 2) with 
two closely approximated posteromedial 
spinose patches, together making up a 
prominent posteromedial lobe; basal parts 
of spinose patches with continuous rows of 
spines, distal parts tapered and with smaller, 
separated spinules. Sternite 6 simple medi- 
ally, expanded into a broad, pale part con- 
nected with a broad, thin-rimmed ring scler- 
ite posteriorly. Epandrium (Fig. 3) uniform- 
ly setose; surstylus with a quadrate, ven- 
trally notched anteroventral lobe, a rounded 
posteroventral lobe with 3 stout bristles, 
and a densely long-setose lateral swelling. 
Cercus (Fig. 4) weakly differentiated from 



epandrium, bare dorsally and medially, long 
setose ventrolaterally, cerci bent into broad 
posterior lobes at middle, not fused to form 
a subanal plate but narrowly contiguous 
ventrally. Subepandrial sclerite very broad, 
quadrate, articulating with inner ventral 
corners of cercus medially and posterior 
dorsal corners of surstyli laterally. Hypan- 
drium with a well developed anterior apo- 
deme; lateral arm fused with epandrium and 
contiguous with surstylus. Postgonite (Fig. 
1) (paramere or gonostylus of authors) 
weakly S-shaped, distal part swollen at 
bend then narrowed at apex. Basiphallus 
simple, rounded; distiphallus simple, nar- 
row and tubular, ending in two dark lobes 
and a small, weakly spinulose membrane. 
Ejaculatory apodeme well developed. 

Female abdomen (Figs. 5-6): Tergite 8 
equal in length to tergite 7, but with tripar- 
tite pigmentation, middle part small and 
pale. Tergite 10 pale anteromedially and 
posteromedially, with a bristle on each half. 
Cercus short, with 2 long, thin apical bris- 
tles and a long, thin dorsal bristle. Sternite 
8 small, subequal in length to sternite 7 but 
less than half as wide, bare on anterior 
third, otherwise setulose and setose. Vagi- 
nal area weakly sclerotised with 2 ring- 
shaped sclerites. Sternite 10 densely setu- 
lose on posterior half, with a marginal row 
of bristles; anterior half pale and bare, with 
a deep, keyhole-shaped anteromedial depig- 
mented area. Spermathecae (Fig. 7) dark, 
acorn-shaped; sclerotised parts of ducts 
very short. 

Holotype.— UNITED STATES. Florida: 
Levy Co., Archer, on rotting fungus on 
sand, 17.xii.l997, S.A. Marshall (6, Uni- 
versity of Guelph). 

Paraty pes. —UNITED STATES. Florida: 
Marion Co., Ocala National Forest, 
29.1.1986, flight intercept trap, R.&M. Mar- 
shall (2$, University of Guelph). 

Xenolimosina phoba Marshall 
(Figs. 8-10) 

Xenolimosina phoba Marshall 1985: 764 
(male only). 



VOLUME 101, NUMBER 2 



273 



Description of female terminalia (Figs. 
8-10). — Tergite 8 completely divided; ter- 
gite 10 uniformly pigmented posteriorly, 
anteromedially pale. Cercus long, thin, with 
long apical, preapical and dorsal bristles. 
Sternite 8 large, longer than sternite 7 and 
over half as wide, almost entirely setulose, 
with 2 transverse rows of bristles. Sternite 
10 with tripartite pigmentation anteriorly, 
middle part bare. Spermathecae (Fig. 8) 
acorn-shaped, sclerotised parts of ducts 
very short. 

New records since 1985. — CANADA. 
Quebec, Old Chelsea, 17.x. 1988, J.R. 
Vockeroth (1 9 , Canadian National Collec- 
tion, Ottawa). Ontario, St. Joseph's Island, 
Hilton Beach, fish entrail baited pan traps 
in hardwood forest, ix-x.l987, J. Swann (8 
6 , University of Guelph). 

Comments. — The first and only female 
of this species was collected by Dr. Richard 
Vockeroth, who collected the male holotype 
at the same locality, and during the same 
month and week, twenty four years earlier. 



Xenolimosina sicula Marshall 
Xenolimosina sicula Marshall 1985: 765. 

New distributional records. — In addition 
to the original records from Ontaiio, Que- 
bec, California and Arkansas, specimens 
have been examined from the following lo- 
calities: CANADA. British Columbia: Car- 
manah Valley. Ontario: Wellington Co., Al- 
goma Co. UNITED STATES. New Hamp- 
shire: Strafford Co. Florida: Leon Co. All 
new records are from October-November. 

Literature Cited 

Marshall, S. A. 1985. The genera Xenolimosina and 
Tenilimosina (Diptera: Sphaeroceridae: Limosi- 
ninae) in North America. Proceedings of the En- 
tomological Society of Washington 87: 759-769. 

Rohacek, J. 1982-3. A monograph and reclassification 
of the previous genus Limosina Macquart (Dip- 
tera, Sphaeroceridae) of Europe. Part I, Beitrage 
zur Entomologie 32: 195-282 and Part 2 Beitrage 
zur Entomologie 33: 3-195. 



PROC. ENTOMOL. SOC. WASH. 
101(2). 1999, pp. 274-284 

LIST OF SPECIES OF NEOTROPICAL MEGALOPTERA (NEUROPTERIDA) 

Atilano Contreras-Ramos 

Institute de Biologia, U.N. A.M., Departamento de Zoologia, Apdo. Postal 70-153, 
04510 Mexico, D. F., Mexico (e-mail: atilano@mail.ibiologia.unam.mx) 



Abstract. — -The 63 known species and subspecies of Neotropical dobsonflies and al- 
derflies (Neuropterida: Megaloptera) are listed. Synonymy, authors, bibliographic refer- 
ences, and distribution of species by country and generally by province or state are in- 
cluded. Six localities, each of specimens thought to belong to new species, are also pro- 
vided. 

Resumen. — Se proporciona una lista de las 63 especies y subespecies conocidas de 
megalopteros (Neuropterida: Megaloptera) neotropicales. Se incluye sinoniinia, autores, 
referencias bibliograficas y la distribucion de las especies por pais y generalmente tambien 
por provincia o estado. Se enumeran seis localidades de ejemplares que se cree pertenecen 
a especies nuevas para la ciencia. 

Key Words: Corydalidae, Sialidae, dobsonflies, fishflies, alderflies, Megaloptera, Noo- 
tropics, taxonomy, species list 



Megaloptera is a relatively primitive en- 
dopterygote (holometabolous) group close- 
ly related to Raphidioptera and Planipennia 
(Neuroptera s. s.). Despite being a small 
group, of approximately 200 (Evans 1972) 
to 300 species (New and Theischinger 
1993) worldwide, the alpha taxonomy of 
the Neotropical megalopteran fauna has 
only recently reached a satisfactory level of 
knowledge. Since the most recent species 
list for the Nootropics (Penny 1977) does 
not include results of subsequent revisions, 
it seems useful to provide a current list of 
the species with references to relevant lit- 
erature (i.e., original species descriptions, 
taxonomic revisions, significant distribution 
data). Species synonymy according to the 
most recent revision is included, as are spe- 
cies distributions by country and province 
or state if the latter were published or could 
be obtained. Ordering of taxa is alphabeti- 
cal. Several records {Corydalus and Platy- 



neuromus) are given as "sp. 1, sp. 2", etc., 
and for these locality data are given. This 
is in the hope that new material may be 
collected and the identity of these possibly 
new species might be established. Follow- 
ing Penny (1977), species of Sialidae aie 
included in Protosialis. However, their ex- 
act generic position is rather uncertain at 
this time, as the group needs revision (e.g., 
Ross [1937] synonymized Protosialis under 
Sialis). In accordance to Cabrera and Wil- 
link's (1980) limits of the Neotropical Re- 
gion, the list includes all species and sub- 
species recorded south of the United 
States-Mexico border. The corydalid spe- 
cies Corydalus cornutus (L.) and Dysmi- 
coherrnes sp. have been collected in Texas, 
close to or at the Mexican border (Contrer- 
as-Ramos 1995b, 1998), so there is the pos- 
sibility they occur in Mexico (Tamaulipas). 
Davis (1903) included Mexico as part of the 
distribution range of Nigronia fasciatus 



VOLUME 101, NUMBER 2 



275 



(Walker). However, the southernmost dis- 
tribution otherwise recorded for this species 
is in Florida (Hazard 1960). 

The 63 valid species and subspecies in 
this list, contrast to the 46 species recorded 
for the Megaloptera of America north of 
Mexico (Evans and Neunzig 1996; 43 spe- 
cies [3 Corydalus spp. missing] in Penny et 
al. 1997). The highest diversity in the Neo- 
tropics lies within Corydalinae (46 spp., 
compared to 9 species and subspecies of 
Chauliodinae and 8 species of Sialidae), 
whereas Sialidae is the most diverse group 
in the Nearctic (24 spp., compared to 18 
species of Chauliodinae and 4 species of 
Corydalinae). Four species ( 1 Chauliodinae 
and 3 Corydalinae) are shared between both 
regions. 

Family Corydalidae 

Subfamily Chauliodinae 
(fishflies) 

Genus Archichauliodes Weele 1909 

1. Archichauliodes chilensis Kimmins. 
Archichauliodes chilensis Kimmins 1954: 

Flint 1973; Penny 1977. 

Distribution. — CHILE: Arauco, Coquim- 
bo, Curico, Malleco, Santiago, Valdivia, 
Valparaiso. 

2. Archicahuliodes pinares Flint. 
Archicahuliodes pinares Flint 1973. 

Distribution. — CHILE: Concepcion. 

Genus Neohermes Banks 1908 

3. Neohermes filicornis (Banks). 
Chauliodes filicornis Banks 1903b. 
Neohermes filicornis (Banks): Weele 1910; 

Flint 1965; Evans 1972; Contreras-Ra- 
mos 1991a, 1997. 

Distribution.— UNITED STATES: Ari- 
zona, California, New Mexico; MEXICO: 
Baja California, Sonora. 

Genus Nothochauliodes Flint 1983 

4. Nothochauliodes penai Flint. 
Nothochauliodes penai Flint 1983. 



Distribution. — CHILE: Maule. 

Genus Protochauliodes Weele 1909 

5. Protochauliodes bullocki Flint. 
Protochauliodes bullocki Flint 1973. 

Distribution. — CHILE: Bio-Bio, Linares, 
Malleco, Nuble. 

6. Protochauliodes cinerascens cinerascens 
(Blanchard). 

Chauliodes cinerascens Blanchard 1851. 
Neohermes cinerascens (Blanchard): Banks 

1908. 
Protochauliodes cinerascens (Blanchard): 

Weele 1910; Kimmins 1954. 
Protochauliodes cinerascens cinerascens 

(Blanchard): FHnt 1973. 

Distribution. — CHILE: Curico, Linares, 
Nuble, O'Higgins, Santiago, Talca. 

7. Protochauliodes cinerascens fumipennis 
Flint. 

Protochauliodes cinerascens fumipennis 
Flint 1973. 

Distribution. — CHILE: Concepcion. 

8. Protochauliodes cinerascens reedi Kim- 
mins. 

Protochauliodes reedi Kimmins 1954. 
Protochauliodes cinerascens reedi Kim- 
mins: Flint 1973. 

Distribution. — CHILE: Valparaiso. 

9. Protochauliodes humeralis (Banks). 
Neohermes humeralis Banks 1908. 
Protochauliodes humeralis (Banks): Weele 

1910; Flint 1973. 

Distribution. — CHILE: Arauco, Bio-Bio, 
Cautin, Concepcion, Malleco, Maule. 

Subfamily Corydalinae 
(dobsonflies) 

Genus Chloronia Banks 1908 

10. Chloronia absona Flint. 

Chloronia absona Flint 1992: Contreras- 
Ramos 1995a. 

Distribution. — COSTA RICA: Alajuela, 
Guanacaste, Limon, San Jose. 



276 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



1 1 . Chloronia antiUiensis Flint. 
Chloronia antiUiensis Flint 1970: Penny 

and Flint 1982; Contreras-Ramos 1995a. 

Distribution. — DOMINICA. 

12. Chloronia banksiana Penny and Flint. 
Chloronia banksiana Penny and Flint 1982: 

Contreras-Ramos 1995a. 
Chloronia bogotana (not Weele, misidenti- 
fication): Banks 1943. 

Distribution. — VENEZUELA: Aragua, 
Carabobo. 

13. Chloronia bogotana Weele. 
Chloronia bogotana Weele 1909: Penny 

and Flint 1982; Flint 1991; Contreras-Ra- 
mos 1995a. 

Distribution.— BOLIVIA: La Paz; CO- 
LOMBIA: Cundinamarca; ECUADOR: Za- 
mora-Chinchipe; PERU: Cuzco, Huanuco. 

14. Chloronia convergens Contreras-Ra- 
mos. 

Chloronia convergens Contreras-Ramos 
1995a. 

Distribution.— ECUADOR: Pichincha. 

15. Chloronia corripiens (Walker). 
Hermes corripiens Walker 1858. 
Neuromus corripiens (Walker): Mac- 

Lachlan 1869. 
Neuromus winthemi Davis 1903. 
Chloronia meridionalis Weele 1909. 
Chloronia winthemi (Davis): Weele 1910. 
Chloronia ocellaris Navas 1934a. 
Chloronia corripiens (Walker): Penny and 

Flint 1982; Contreras-Ramos 1995a. 

Distribution. — BRAZIL: Espirito Santo, 
Minas Gerais, Rio de Janeiro, Santa Catar- 
ina, Sao Paulo. 

16. Chloronia gloriosoi Penny and Flint. 
Chloronia gloriosoi Penny and Flint 1982: 

Flint 1992; Contreras-Ramos 1995a. 

Distribution. — COSTA RICA: San Jose; 
PANAMA: Chiriqui. 

17. Chloronia hieroglyphica (Rambur). 
Neuromus hieroglyphica Rambur 1842. 



Hermes hieroglyphicus (Rambur): Walker 
1853. 

Corydalis hieroglyphicus (Rambur): Hagen 
1861. 

Chloronia hieroglyphica (Rambur): Banks 
1908; Penny and Flint 1982; Contreras- 
Ramos 1995a. 

Distribution. — BRAZIL: Amazonas, 
Para; FRENCH GUIANA; GUYANA; 
PERU: Loreto, Madre de Dios. 

18. Chloronia me.xicana Stitz. 
Chloronia me.xicana Stitz 1914: Penny and 

Flint 1982; Flint 1992; Contreras-Ramos 
1995a, 1997. 

Distribution. — COSTA RICA: Alajuela, 
Guanacaste, Heredia; GUATEMALA: Alta 
Verapaz, Suchitepequez; MEXICO: Chia- 
pas, Morelos, San Luis Potosi, Tamaulipas, 
Veracruz. 

19. Chloronia mirifica Navas. 
Chloronia mirifica Navas 1925a: Penny and 

Flint 1982; Flint 1992; Contreras-Ramos 
1995a, 1997. 
Chloronia hieratica Navas 1928a. 

Distribution.— COLOMBIA: Meta; 
COSTA RICA: Alajuela, Cartago, Guana- 
caste, Heredia, Limon, Puntarenas, San 
Jose; ECUADOR: Napo, Pichincha; GUA- 
TEMALA: Alta Verapaz, Izabal, Solola; 
MEXICO: Oaxaca, Veracruz; PANAMA: 
Chiriqui, Colon; PERU: Huanuco. 

20. Chloronia osae Flint. 

Chloronia osae Flint 1992: Contreras-Ra- 
mos 1995a. 

Distribution. — COSTA RICA: Puntaren- 
as. 

21. Chloronia pallida (Davis). 
Neuromus pallidus Davis 1903. 
Chloronia pallidus (Davis): Penny 1977. 
Chloronia pallida (Davis): Penny and Flint 

1982; Contreras-Ramos 1995a, 1997. 

Distribution. — MEXICO: Chihuahua, 
Guerrero, Jalisco, Michoacan, Morelos, 
Nayarit. 



VOLUME 101, NUMBER 2 



111 



11. Chloronia plaumanni Penny and Flint. 
Chloronia plaumanni Penny and Flint 
1982: Contreras-Ramos 1995a. 

Distribution. — BRAZIL: Rio Grande do 
Sul, Santa Catarina. 

23. Chloronia zacapa Contreras-Ramos. 
Chloronia zacapa Contreras-Ramos 1995a. 



Distribution. 
Zacapa. 



-GUATEMALA: Izabal, 



Genus Corydalus Latreille 1802 

24. Corydalus affinis Burmeister. 
Corydalus affinis Burmeister 1839: Weele 

1910 (in part); Penny 1977; Contreras- 
Ramos 1998. 

Corydalus nubilus (not Erichson, misiden- 
tification): Weele 1910; Banks 1943; 
Penny 1982. 

Corydalus sp.: Glorioso 1981. 

Corydalus spec, nov.: Geijskes 1984. 

Distribution.— ARGENTINA: Chaco, 
Misiones; BOLIVIA: Beni; BRAZIL: Acre, 
Amapa, Amazonas, Mato Grosso, Para, 
Rondonia, Roraima, Sao Paulo; COLOM- 
BIA: Antioquia, Boyaca, Choco, Cundina- 
marca, Tolima; ECUADOR: Napo, Pichin- 
cha, Sucumbi'os; FRENCH GUIANA; 
GUYANA; PARAGUAY; PERU: Cuzco, 
Loreto, Madre de Dios; VENEZUELA: 
Guarico. 

25. Corydalus amazonas Contreras-Ramos. 
Corydalus amazonas Contreras-Ramos 

1998. 

Distribution. — BRAZIL: Amazonas, 
Rondonia. 

26. Corydalus armatus Hagen. 
Corydalis annata Hagen 1861. 
Corydalis armatus Hagen 1861. 
Corydalus armatus Hagen 1861: Weele 

1910 (in part); Stitz 1914 (in part); Navas 
1920a, 1934a, 1935; Penny 1977 (in 
part), 1982 (in part); Glorioso 1981 (in 
part); Contreras-Ramos 1998. 
Corydalis annata n. sp.: Davis 1903 (in 
part). 



Corydalus quadrispinosus Stitz 1914. 
Corydalus peruvianus (not Davis, misiden- 
tification): Banks 1943 (in part). 

Distribution.— ARGENTINA ; Catamar- 
ca, Jujuy, Salta, Tucuman; BOLIVIA: Chu- 
quisaca, Cochabamba, La Paz, Santa Cruz; 
COLOMBIA: Boyaca, Cundinamarca, Val- 
le del Cauca; ECUADOR: Bolivar, Chim- 
borazo. El Oro, Esmeraldas, Imbabura, 
Loja, Los Rios, Napo, Pichincha, Tungura- 
hua, Zamora Chinchipe; PERU: Ayacucho, 
Cuzco, Huanuco, Juni'n, Lima, Pasco; 
VENEZUELA: Aragua, Merida, Tachira. 

27. Corydalus arpi Navas. 

Corydalus arpi Navas 1936: Penny 1977, 
1982; Contreras-Ramos 1993, 1998. 

Distribution. — BRAZIL: Amazonas, 
Rondonia; VENEZUELA: Territorio Fed- 
eral Amazonas. 

28. Corydalus australis Contreras-Ramos. 
Corydalus australis Contreras-Ramos 1998. 
Corydalus affinis (not Burmeister, misiden- 

tification): Weele 1910 (in part); Penny 
1977 (in part); Glorioso 1981. 

Distribution. — ARGENTINA: Misiones; 
BRAZIL: Minas Gerais, Rio Grande do 
Sul, Santa Catarina; URUGUAY: Artigas. 

29. Corydalus batesii MacLachlan. 
Corydalus batesii MacLachlan 1868: Davis 

1903; Geijskes 1984; Contreras-Ramos 
1998. 
Corydalus batesi MacLachlan: Weele 1910; 
Stitz 1914 (in part); Kimmins 1970; Pen- 
ny 1977, 1982; Glorioso 1981. 

Distribution. — BOLIVIA: Cochabamba, 
Santa Cruz; BRAZIL: Amazonas, Para 
COLOMBIA: Antioquia; ECUADOR 
Napo; FRENCH GUIANA; GUYANA 
SURINAME; PERU: Madre de Dios 
VENEZUELA: Territorio Federal Amazon- 
as. 

30. Corydalus bidenticulatus Contreras-Ra- 
mos. 

Corydalus bidenticulatus Contreras-Ramos 
1998. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Corydalus lutea (not Hagen, misidentifica- 

tion): Glorioso 1981 (in part). 
Corydalus sp. B: Contreras-Ramos 1997. 

Distribution.— UNITED STATES: Ari- 
zona; MEXICO: Colima, Guerrero, Jalisco, 
Michoacan, Morelos, Nayarit, Oaxaca, Sin- 
aloa, Sonora. 

31. Corydalus cephalotes Rambur. 
Corydalus cephalotes Rambur 1842: Weele 

1910 (in part); Stitz 1914 (in part); Penny 
1977 (in part), 1982 (in part); Contreras- 
Ramos 1998. 

Corydalis affinis (not Burmeister, misiden- 
tification): Walker 1853; Hagen 1861. 

Neuromus cephalotes (Rambur): Davis 
1903 (in part). 

Corydalus intricatus Navas 1921: Penny 
1977. 

Distribution. — BRAZIL: Rio de Janeiro. 

32. Corydalus clauseni Contreras-Ramos. 
Corydalus clauseni Contreras-Ramos 1998. 

Distribution.— COLOMBIA: Valle del 
Cauca; COSTA RICA: Heredia; ECUA- 
DOR: Caiiar, Loja, Napo, Pichincha, Tun- 
gurahua. 

33. Corydalus colombianus Contreras-Ra- 
mos. 

Corydalus colombianus Contreras-Ramos 
1998. 

Corydalus ecuadorianus (not Banks, mis- 
identification): Glorioso 1981 (in part). 

Distribution.— COLOMBIA: Valle del 
Cauca. 

34. Corydalus diasi Navas. 

Corydalus diasi Navas 1915: Penny 1977; 

Contreras-Ramos 1998. 
Corydalus finoti Navas 1921: Penny 1977. 

Distribution. — ARGENTINA: Misiones, 
BRAZIL: Bahia, Ceara, Goias, Minas Ger- 
ais, Rio Grande do Sul, Sao Paulo; PAR- 
AGUAY. 

35. Corydalus ecuadorianus Banks. 
Corydalus ecuadorianus Banks 1948: Pen- 



ny 1977; Glorioso 1981 (in part); Con- 
treras-Ramos 1998. 



Distribution, 
gurahua. 



-ECUADOR: Napo, Tun- 



36. Corydalus flavicomis Stitz. 
Corydalus armatus flavicomis Stitz 1914: 

Penny 1977. 
Corydalus nevermanni Navas 1934b: Penny 

1977; Banks 1943. 
Corydalus camposi Navas 1935: Penny 

1977. 
Corydalus armatus (not Hagen, misidenti- 

fication): Banks 1943 (in part). 
Corydalus peruvianus (not Davis, misiden- 

tification): Banks 1943 (in part). 
Corydalus flavicomis Stitz: Glorioso 1981; 

Contreras-Ramos 1998. 

Distribution. — COLOMBIA: Antioquia, 
Meta, Santander; COSTA RICA: Alajuela, 
Cartago, Guanacaste, Heredia, Limon, Pun- 
tarenas, San Jose; ECUADOR: Guayas, Es- 
meraldas, Loja, Los Rios, Napo, Pastaza, 
Pichincha, Tungurahua; EL SALVADOR 
GUATEMALA; HONDURAS; PANAMA 
Chiriqui; PERU: Huanuco, Junin, Pasco 
VENEZUELA: Aragua, Bolivar, Lara, Me- 
rida, Portuguesa, Zulia. 

37. Corydalus flinti Contreras-Ramos. 
Corydalus flinti Contreras-Ramos 1998. 

Distribution. — VENEZUELA: Territorio 
Federal Amazonas. 

38. Co/ydalus hecate MacLachlan. 
Corydalis hecate MacLachlan 1866: Kim- 

mins 1970. 

Neuromus cephalotes (not Rambur, mis- 
identification): Davis 1903 (in part). 

Corydalus cephalotes (not Rambur, mis- 
identification): Weele 1910 (in part); Stitz 
1914 (in part); Penny 1977 (in part), 
1982 (in part); Glorioso 1981 (in part). 

Corydalus raymundoi Navas 1920b. 

Corydalus sallei Navas 1920b. 

Corydalus hecate MacLachlan: Contreras- 
Ramos 1998. 



VOLUME 101. NUMBER 2 



279 



Distribution. — BRAZIL: Distrito Feder- 
al, Espirito Santo, Minas Gerais, Sao Paulo; 
PERU; VENEZUELA: Distrito Federal. 

39. Corydalus holzenthali Contreras-Ra- 
mos. 

Corydalus holzenthali Contreras-Ramos 
1998. 

Distribution. — BOLIVIA: Cochabamba, 
La Paz; PERU: Pasco. 

40. Corydalus ignotus Contreras-Ramos. 
Corydalus ignotus Contreras-Ramos 1998. 

Distribution.— FRENCH GUIANA. 

41. Corydalus imperiosus Contreras-Ra- 
mos. 

Corydalus imperiosus Contreras-Ramos 

1998. 
Corydalus tridentatus (not Stitz, misidenti- 

fication): Glorioso 1981 (in part). 

Distribution. — ARGENTINA: Misiones. 

42. Corydalus longicornis Contreras-Ra- 
mos. 

Corydalus longicornis Contreras-Ramos 
1998. 

Distribution.— ARGENTINA: Catamar- 
ca, Salta; BOLIVIA: Chuquisaca, Cocha- 
bamba, Santa Cruz; ECUADOR: Zamora 
Chinchipe. 

43. Corydalus luteus Hagen. 

Corydalis lutea Hagen 1861: Davis 1903 
(in part); Penny 1977 (as nomen nudum). 

Corydalus luteus Hagen: Weele 1910 (as ju- 
nior synonym of C cornutus [L.]); Con- 
treras-Ramos 1997, 1998. 

Corydalus lutea Hagen: Glorioso 1981 (in 
part). 

Corydalis crassicornis MacLachlan 1868: 
Davis 1903; Banks 1907; Kimmins 1970. 

Corydalus crassicornis (MacLachlan): 
Weele 1910 (as junior synonym of C. 
cornutus [L.]). 

Corydalis inamabilis MacLachlan 1868: 
Davis 1903; Banks 1907; Kimmins 1970. 

Corydalus inamabilis (MacLachlan): Weele 
1910 (as junior synonym of C cornutus 
[L.]). 



Corydalus armatus laevicornis Stitz 1914: 
Penny 1977. 

Distribution. — BELIZE; COSTA RICA: 
Alajuela, Guanacaste, Heredia, Limdn, 
Puntarenas, San Jose; EL SALVADOR; 
UNITED STATES: Texas; GUATEMALA: 
Alta Verapaz, Chiquimula, Escuintla, Gua- 
temala, Sacatepequez, San Marcos, Santa 
Rosa, Suchitepequez, Zacapa; HONDU- 
RAS; MEXICO: Chiapas, Coahuila, Hidal- 
go, Nuevo Leon, Oaxaca, Queretaro, San 
Luis Potosi, Tabasco, Tamaulipas, Vera- 
cruz; NICARAGUA; PANAMA: Zona del 
Canal, Chiriqui, Colon. 

44. Corydalus magnus Contreras-Ramos. 
Corydalus magnus Contreras-Ramos 1998. 
Corydalus lutea (not Hagen, misidentifica- 

tion): Glorioso 1981 (in part). 
Corydalus sp. M: Contreras-Ramos 1997. 

Distribution. — COSTA RICA: Alajuela, 
Guanacaste, Puntarenas; EL SALVADOR; 
GUATEMALA: Alta Verapaz, Baja Vera- 
paz, Suchitepequez; MEXICO: Chiapas, 
Puebla, San Luis Potosi, Veracruz. 

45. Corydalus neblinensis Contreras-Ra- 
mos. 

Corydalus neblinensis Contreras-Ramos 
1998. 

Distribution. — VENEZUELA: Territorio 
Federal Amazonas. 

46. Corydalus nubilus Erichson. 
Corydalis nubila Erichson 1848: Hagen 

1861 (in part); Davis 1903 (in part). 

Corydalus nubilus Erichson: Stitz 1914; 
Penny 1977 (in part); Glorioso 1981; Ge- 
ijskes 1984; Contreras-Ramos 1998. 

Corydalus nevermanni (not Navas, mis- 
identification): Penny 1982. 

Corydalus titschacki Navas 1928b: Penny 
1977. 

Distribution. — BRAZIL: Amazonas, 
Para, Roraima; FRENCH GUIANA; GUY- 
ANA; VENEZUELA: Territorio Federal 
Amazonas. 

47. Corydalus parx'us Stitz. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Corydalus parvus Stitz 1914: Penny 1977; 

Contreras-Ramos 1998. 
Corydalus annatus (not Hagen, misidenti- 

fication): Glorioso 1981 (in part). 

Distribution.— ECUADOR: Pastaza, Za- 
mora Chinchipe; PERU: Cuzco, Huanuco, 
Pasco. 

48. Corydalus peruvianus Davis. 
Corydalis peruviana Davis 1903. 
Corydalus armatus (not Hagen, misidenti- 

fication): Weele 1910 (in part); Stitz 1914 
(in part); Penny 1977 (in part), 1982 (in 
part); Glorioso 1981 (in part). 

Corydalis crassicornis (not MacLachlan, 
misidentification): Banks 1914. 

Corydalus primitivus fera Navas 1927: Pen- 
ny 1977. 

Corydalus peruvianus Davis: Banks 1943 
(in part); Contreras-Ramos 1997, 1998. 

Distribution. — ARGENTINA: Jujuy, 
Salta, Tucuman; BOLIVIA: Chuquisaca, 
Cochabamba, La Paz, Tarija; COLOMBIA: 
Antioquia, Cundinamarca, Meta, Putumayo, 
Valle del Cauca; COSTA RICA: Alajuela, 
Cartago, Guanacaste, Heredia, Limon, Pun- 
tarenas, San Jose; ECUADOR: Cotopaxi, 
Guayas, Los Rios, Morona Santiago, Napo, 
Pastaza; Pichincha; Sucumbios; GUATE- 
MALA: Escuintla, Izabal, Quiche, San 
Marcos; MEXICO: Chiapas, Oaxaca, Pueb- 
la. Tabasco, Veracruz; PANAMA: Bocas 
del Toro, Chiriqui, Colon, Darien; PERU: 
Amazonas, Huanuco, La Libertad, Pasco; 
VENEZUELA: Aragua, Barinas, Distrito 
Federal, Merida. 

49. Corydalus primitivus Weele. 
Corydalus primitivus Weele 1909: Weele 

1910 (in part); Stitz 1914; Navas 1929; 
Penny 1977, 1982 (in part); Glorioso 
1981 (in part), Contreras-Ramos 1998. 

Distribution. — ARGENTINA: Catamar- 
ca, Jujuy, Salta, Tucuman; BOLIVIA: Santa 
Cruz. 

50. Corydalus tesselatus Stitz. 
Corydalus batesi tesselatus Stitz 1914. 
Corydalis nubila (not Erichson, misidenti- 



fication): Hagen 1861 (in part); Davis 
1903 (in part); Banks 1943. 

Corydalus bolivari Banks 1943: Glorioso 
1981; Contreras-Ramos 1993. 

Corydalus tesselatus Stitz: Penny 1977 (in- 
correct locality); Contreras-Ramos 1998. 

Distribution.— COLOMBIA; VENE- 
ZUELA: Aragua, Distrito Federal, Merida, 
Tachira. 

51. Corydalus texanus Banks. 
Corydalis te.xana Banks 1903a: Penny 1977 

(as junior synonym of C. cornutus [L.]). 

Corydalis cognata (not Hagen, misidentifi- 
cation): Banks 1892, 1903b, 1907; Davis 
1903; Chandler 1956. 

Neuromus pallidus (not Davis): Davis 1903 
(mislabeled photograph). 

Corydalus pallidus (not Davis): Weele 1910 
(invalid combination based on mislabeled 
photograph; as junior synonym of C cor- 
nutus [L.]). 

Corydalus cognatus (not Hagen, misiden- 
tification): Weele 1910 (in part; as junior 
synonym of C. cornutus [L.]); Evans 
1972 (in part). 

Corydalus texanus Banks: Weele 1910 (as 
junior synonym of C. cornutus [L.]); 
Contreras-Ramos 1997, 1998. 

Corydalus cornutus (not Linnaeus, mis- 
identification): Stitz 1914 (in part); Glo- 
rioso 1981 (in part); Hermann and Davis 
1991. 

Cory^dalus similis Stitz 1914: Banks 1943 
(misidentification); Penny 1977 (in part). 

Corydalus constellatus Navas 1934b: Pen- 
ny 1977. 

Distribution.— UNITED STATES: Ari- 
zona, California, Colorado, Nevada, New 
Mexico, Texas, Utah; GUATEMALA: Baja 
Verapaz, Chiquimula; MEXICO: Baja Cal- 
ifornia, Baja CalifoiTiia Sur, Chiapas, Chi- 
huahua, Colima, Distrito Federal, Guerrero, 
Jalisco, Michoacan, Morelos, Nayarit, Oa- 
xaca, Puebla, Queretaro, Sinaloa, Sonora, 
Veracruz. 

52. Corydalus tridentatus Stitz. 
Corydalus tridentatus Stitz 1914: Penny 



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281 



1977; Glorioso 1981 (in part); Contreras- 
Ramos 1998. 
Corydahis tridentatits nigripes Stitz 1914. 

Distribution. — BRAZIL: Espirito Santo. 
Parana, Rio Grande do Sul. 

Corydalus sp. 1. 

Coi-ydalus sp. 1 (near C. affinis Burmeis- 
ter): Contreras-Ramos 1998. 

Distribution. — BRAZIL: Sao Paulo: Sao 
Jose dos Campos. 

Corydalus sp. 2 

Contains tesselatus (not Stitz, misidenti- 
fication): Banks 1943. 

Coiydalus ecuadorianus (not Banks, mis- 
identification): Glorioso 1981 (in part). 

Coiydalus sp. 2 (near C. ecuadorianus 
Banks): Contreras-Ramos 1998. 

Distribution. — COLOMBIA: [Cundina- 
marca?]: St. [San] Antonio, 2000 m. 

Corydalus sp. 3. 

Corydalus sp. 3 (near C. nubilus Erichson): 
Contreras-Ramos 1998. 

Distribution.— VENEZUELA: Bolivar: 
Ciudad Bolivar 

Corydalus sp. 4. 

Corydalus sp. 4 (near C tesselatus Stitz): 
Contreras-Ramos 1998. 

Distribution. — ECUADOR: Sucumbios: 
El Reventador, [aprox. 900 m]. 

Corydalus sp. 5. 

Corydalus sp. 5 (near C colombianus Con- 
treras-Ramos): Contreras-Ramos 1998. 

Distribution. — ECUADOR: Pichincha: 
Palmeras. 

Genus Platyneuromus Weele 1909 

53. Platyneuromus honduranus Navas. 

Platyneuromus soror hondurana Navas 
1928b. 

Platyneuromus auritus Kimmins 1928. 

Platyneuromus honduranus Navas: Glorio- 
so and Flint 1984; Contreras-Ramos 
1997. 



Distribution. — GUATEMALA: Alta Ver- 
apaz. El Peten, Izabal; HONDURAS: At- 
lantida, Cortes; MEXICO: Chiapas. 

54. Platyneuromus reflexus Glorioso and 
Flint. 

Platyneuromus reflexus Glorioso and Flint 
1984: Contreras-Ramos 1997. 

Distribution.— GUATEMALA: Alta Ver- 
apaz; MEXICO: Chiapas. 

55. Platyneuromus soror (Hagen). 
Corydalis soror Hagen 1861. 
Neuromus soror (Hagen): Davis 1903. 
Neuromus (Chloronia) soror (Hagen): 

Banks 1908. 

Platyneuromus soror (Hagen): Weele 1909, 
1910; Stitz 1914; Penny 1977; Glorioso 
1981; Glorioso and Flint 1984; Contrer- 
as-Ramos 1991b, 1997; Contreras-Ramos 
and Harris 1998. 

Doeringia christel Navas 1925b. 

Distribution.— COSTA RICA: Alajuela, 
Cartago, Guanacaste, Heredia, Puntarenas, 
San Jose; MEXICO: Chiapas, Distrito Fed- 
eral, Hidalgo, Estado de Mexico, Nuevo 
Leon, Oaxaca, Puebla, Queretaro, San Luis 
Potosi, Tamaulipas, Veracruz; PANAMA: 
Chiriqui. 

Platyneuromus sp. 

Platyneuromus larval form A: Contreras- 
Ramos and Harris 1998. 

Distribution. — MEXICO: Guerrero: 56 
km NE Atoyac on road to Puerto del Gallo, 
17.417°N, 100.217°W, 1372 m; Sinaloa: 
4.83 km W Palmito. 

Family Sialidae 
(alderflies) 

Genus Protosialis Weele 1909 

56. Protosialis bifasciata (Hagen). 
Sialis bifasciata Hagen 1861: Davis 1903. 
Protosialis bifasciata (Hagen): Weele 1910; 

Penny 1977. 

Distribution. — CUBA. 

57. Protosialis bimaculata Banks. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Protosialis bimacitlata Banks 1920: Penny 
1977, 1981 [1982]. 

Distribution. — BOLIVIA: La Paz. 

58. Protosialis brasiliensis Navas. 
Protosialis brasiliensis Navas 1936: Penny 

1977, 1981 [1982]. 

Distribution. — BRAZIL: Sao Paulo. 

59. Protosialis chilensis (MacLachlan). 
Sialis chilensis MacLachlan 1870: Davis 

1903; Flint 1973. 
Protosialis chilensis (MacLachlan): Weele 
1910; Penny 1977. 

Distribution. — CHILE: Arauco, Concep- 
cion, Llanquihue, Malleco, O'Higgins, Tal- 
ca, Valdivia. 

60. Protosialis flammata Penny. 
Protosialis flamatta Penny 1981 [1982]. 

Distribution. — BRAZIL: Amazonas. 

61. Protosialis flavicollis (Enderlein). 
Sialis flavicollis Enderlein 1910. 
Protosialis flavicollis (Enderlein): Penny 

1977, 1981 [1982]. 

Distribution.— COLOMBIA: Tolima. 

62. Protosialis me.xicana (Banks). 

Sialis me.xicana Banks 1901: Henry et al. 

1992. 
Protosialis mexicana (Banks): Weele 1910; 

Penny 1977, E. D. Evans {in litt.)\ Con- 

treras-Ramos 1991a. 

Distribution. — MEXICO: Chiapas, Vera- 
cruz; PANAMA. 

63. Protosialis nubila Navas. 
Protosialis nubila Navas 1933: Penny 

1977, 1981 [1982]. 

Distribution. — BRAZIL: Matto Grosso? 

Acknowledgments 

Many of the references cited in this paper 
were obtained during my Ph.D. research at 
the Department of Entomology, University 
of Minnesota, St. Paul (UMSP). Thanks to 
Drs. Ralph W. Holzenthal and Philip J. 



Clausen for all their support while I was a 
student at UMSP. Thanks as well to the Di- 
reccion General de Asuntos del Personal 
Academico of UNAM for giving me a doc- 
toral fellowship, to Fernando Muiioz-Que- 
sada (UMSP) who kindly helped obtain 
several missing references, and to two 
anonymous reviewers for improving the 
quality of the manuscript. Several new rec- 
ords from Mexico were obtained through 
project K022 funded by CONABIO. 

Literature Cited 

Banks, N. 1892. A synopsis, catalogue, and bibliog- 
raphy of the neuropteroid insects of temperate 
North America. Transactions of the American En- 
tomological Society 19: 327-373. 

. 1901. A list of neuropteroid insects from 

Mexico. Transactions of the American Entomo- 
logical Society 27: 361-371. 

. 1903a. Some new neuropteroid insects. Jour- 
nal of the New York Entomological Society 1 1 : 
236-243. 

. 1903b. Neuropteroid insects from Arizona. 

Proceedings of the Entomological Society of 
Washington 5: 237-245. 

. 1907. Catalogue of the neuropteroid insects 

(except Odonata) of the United States. American 
Entomological Society, Philadelphia, Pennsylva- 
nia. 53 pp. 

. 1908. On the classification of the Corydalinae, 

with description of a new species. Proceedings of 
the Entomological Society of Washington 10: 27— 
30. 

. 1914. Neuroptera and Trichoptera from Costa 

Rica. Entomological News 25: 149-150. 

. 1920. New neuropteroid insects. Bulletin of 

the Museum of Comparative Zoology, Harvard 
University 64: 299-362. 

. 1943. Neuroptera of northern South America. 



Boleti'n de Entomologia Venezolana 2: 59-66. 
. 1948. A new species of Coiydcilus (Neurop- 



tera). Psyche 55: 82-83. 

Blanchard, E. 1851. Neuropteros, pp. 85-142 In Gay, 
C Historia Fisica y Poli'tica de Chile, Zoologi'a, 
Volumen 6. 

Burmeister, H. 1839. Handbuch der entomologie (Ber- 
lin), Vol. 2. parts 2-3: 397-1050. 

Cabrera, A. and A. Willink. 1980. Biogeografi'a de 
America Latina. Organizacion de los Estados 
Americanos, Monografias Cienti'ficas, Ser. Biolo- 
gi'a 13: 1-122. 

Chandler, H. P 1956. Megaloptera. pp. 229-233. In 
Usinger, R. L., ed.. Aquatic Insects of California. 
University of California Press, Berkeley, Califor- 
nia. 



VOLUME 101. NUMBER 2 



283 



Contreras-Ramos. A. 1991a. Distribution of the Me- 
galoptera in Mexico. Bulletin of the North Amer- 
ican Benthological Society 8(1): 70. 

. 1991b. The first-instar larvae and pupae of 

Platyneuromus and Corydalus (Megaloptera: Cor- 
ydalidae: Corydalinae). Bulletin of the North 
American Benthological Society 8(1): 163. 

. 1993. Four unique dobsonflies (Megaloptera: 

Corydalidae: Corydalinae) from South America. 
Bulletin of the North American Benthological So- 
ciety 10(1): 130-131. 

. 1 995a. New species of Chloronia from Ec- 
uador and Guatemala, with a key to the species in 
the genus (Megaloptera: Corydalidae). Journal of 
the North American Benthological Society 14: 
108-114. 

. 1995b. A remarkable range extension for the 

fishfly genus Dysmicohermes (Megaloptera: Cor- 
ydalidae). Entomological News 106: 123-126. 

. 1997. Clave para la identificacion de los Me- 
galoptera (Neuropterida) de Mexico. Dugesiana 
4(2): 51-61. 

. 1998. Systematics of the dobsonfly genus 

Cotydalus (Megaloptera: Corydalidae). Thomas 
Say Publications, Entomological Society of Amer- 
ica. 

Contreras-Ramos, A. and S. C. Harris. 1998. The im- 
mature stages of Platyneuromus (Corydalidae), 
with a key to the genera of larval Megaloptera of 
Mexico. Journal of the North American Bentho- 
logical Society 17: 489-517. 

Davis, K. C. 1903. Aquatic Insects in New York State. 
Part 7. Sialididae [sic] of North and South Amer- 
ica. New York State Museum, Bulletin 68: 442- 
486, 499, 2 pis. 

Enderlein, G. 1910. Eine neue Sialis aus Columbien. 
Stettiner entomologische Zeitung 1910: 380-381. 

Erichson, W. E 1848. Insekten, pp. 583-586 //; M. R. 
Schomburgk's Reisen in British-Guiana in den 
Jahren 1840-1884, Vol. 3. J. J. Weber, Leipzig. 

Evans, E. D. 1972. A study of the Megaloptera of the 
Pacific coastal region of the United States. Ph.D. 
dissertation, Oregon State University, Corvallis. 

Evans, E. D. and H. H. Neunzig. 1996. Megaloptera 
and aquatic Neuroptera, pp. 298-308. //; Merritt, 
R. W. and K. W. Cummins, eds.. An Introduction 
to the Aquatic Insects of North America, 3rd edi- 
tion. Kendall/Hunt Publishing Company, Du- 
buque, Iowa. 

Flint, O. S., Jr. 1965. The genus Neohermes (Mega- 
loptera: Corydalidae). Psyche 72: 255-263. 

. 1970. The Megaloptera of Dominica. Pro- 
ceedings of the Entomological Society of Wash- 
ington 72: 240-242. 

. 1973. The Megaloptera of Chile (Neuroptera). 



tera: Corydalidae). Entomological News 94: 15- 
17. 

. 1 99 1 . On the identity of Chloronia bogatana 

[sic] Weele (Neuropterida: Megaloptera: Corydal- 
idae). Proceedings of the Entomological Society 
of Washington 93: 489-494. 

. 1992. A review of the genus Chloronia in 

Costa Rica, with the description of two new spe- 
cies (Neuropterida: Megaloptera: Corydalidae). 
Proceedings of the Biological Society of Wash- 
ington 105: 801-809. 

Geijskes, D. C. 1984. Notes on Megaloptera from the 
Guyanas, S. Am., pp. 79-84. In Gepp, J., H. As- 
pock, and H. Holzel, eds.. Progress in World's 
Neuropterology; Proceedings of the 1st Interna- 
tional Symposium on Neuropterology. Graz, Aus- 
tria. 

Glorioso, M. J. 1981. Systematics of the dobsonfly 
subfamily Corydalinae (Megaloptera: Corydali- 
dae). Systematic Entomology 6: 253-290. 

Glorioso. M. J. and O. S. Flint, Jr. 1984. A review of 
the genus Platyneuromus (Insecta: Neuroptera: 
Corydalidae). Proceedings of the Biological So- 
ciety of Washington 97: 601-614. 

Hagen, H. 1861. Synopsis of the Neuroptera of North 
America with a list of the South American spe- 
cies. Smithsonian Miscellaneous Collections 4(1): 
XX + 1-347. 

Hazard, E. I. 1960. A revision of the genera Chauli- 
odes and Nigronia (Megaloptera: Corydalidae). 
M.S. thesis. The Ohio State University. 

Henry, C. S., N. D. Penny, and P A. Adams. 1992. 
The neuropteroid orders of Central America (Neu- 
roptera and Megaloptera), pp. 432-458. //; Quin- 
tero, D., and A. Aiello, eds.. Insects of Panama 
and Mesoamerica. Oxford University Press, Ox- 
ford. 

Herrmann, S. J. and H. L. Davis. 1991. Distribution 
records of Coiydalus cornutus (Megaloptera: Cor- 
ydalidae) in Colorado. Entomological News 102: 
25-30. 

Kimmins, D. E. 1928. New and little known Neurop- 
tera of Central America. Eos 4: 363-370. 

. 1954. A new genus and some new species of 

the Chauliodini (Megaloptera). Bulletin of the 
British Museum (Natural History), Entomology 3: 
417-444. 

. 1970. A list of the type-specimens of Plecop- 



Revista Chilena de Entomologia 7: 31-45. 

. 1983. Nothochauliodes penai, a new genus 

and species of Megaloptera from Chile (Neurop- 



tera and Megaloptera in the British Museum (Nat- 
ural History). Bulletin of the British Museum 
(Natural History), Entomology 24(8): 337-361. 

Latreille, P. A. 1802. Histoire Naturelle, Generale et 
Particuliere, des Crustaces et des Insectes. Tome 
troisieme (Vol. 3). F Dufart, Paris. 

MacLachlan, R. 1866. Description of a new neurop- 
terous insect belonging to the genus Corydalis, 
Latreille. Journal of Entomology (London) 2: 
499-500, 1 pi. 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



. 1868. New genera and species, &c., of neu- 

ropterous insects; and a revision of Mr. F. Wallcer's 
British Museum catalogue of Neuroptera, part ii 
(1853), as far as the end of the genus Mynneleoti. 
Journal of the Linnean Society (Zoology) 9: 230- 
281, 1 pi. 

. 1869. Considerations on the neuropterous ge- 
nus Chaiiliodes and its allies; with notes and de- 
scriptions. The Annals and Magazine of Natural 
History 4(4): 35-46. 

. 1870. On the occurrence of the neuropterous 

genus Sialis in Chili. Entomologist's Monthly 
Magazine 7: 145-146. 

Navas, L. 1915. Neuropteros sudamericanos. Segunda 
serie. Broteria (Serie Zoologica) 13: 5—13. 

. 1920a. Insectos sudamericanos (2a serie). An- 

ales de la Sociedad Cienti'fica Argentina 90: 44- 
51. 

. 1920b. Insectos de America. Boleti'n de la So- 
ciedad Entomologica de Espafia 3: 90—99. 

. 1921. Insectos americanos nuevos o criticos. 

1921. Broteria (Serie Zoologica) 19: 113-124. 

. 1925a. Insectos exoticos nuevos o poco con- 



ocidos, Segunda Serie. Memorias de la Real Ac- 
ademia de Ciencias y Artes de Barcelona, serie 3, 
19: 181-200. 

. 1925b. Neuropteren, Megalopteren, Plecop- 

teren und Trichopteren aus dem Deutschen Ento- 
mologischen Institiit (Berlin-Dahlem). I. serie. — 
Entomologische Mitteilungen 14: 205-212. 

. 1927. Insectos del Museo de Pan's (4a serie). 

Broteria (Serie Zoologica) 24: 5-33. 

. 1928a. Insectos del Museo de Estocolmo. Re- 
vista de la Real Academia de Ciencias Exactas, 
Fisicas y Naturales de Madrid 24: 1-12. 

. 1928b. Insectos del Museo de Hamburgo (pri- 

mera serie). Boleti'n de la Sociedad Entomologica 
de Espafia 11: 59-67. 

. 1929. Insectos de la Argentina (quinta serie). 

Revista de la Sociedad Entomologica Argentina 
2: 219-225. 

. 1933. Decadas de insectos nuevos. Broteria 2: 

34-44, 101-110. 

. 1934a. Insectos suramericanos (octava serie). 

Revista de la Academia de Ciencias de Madrid 31 : 
9-28. 

. 1934b. Insectos del Mu.seo de Hamburgo (2a 



serie). Memorias de la Academia de Ciencias y 

Artes de Barcelona 23: 499-508. 
. 1935. Insectos suramericanos (decima serie). 

Revista de la Academia de Ciencias Exactas, Fi's- 

ico-Qui'micas y Naturales de Madrid 32: 360-375. 
. 1936. Insectos del Brasil (5a. serie). Revista 



do Museu Paulista 20: 722-734. 

New, T. R. and G. Theischinger. 1993. Megaloptera 
(Alderflies, Dobsonflies). Handbuch der Zoologie, 
Vol. 4 (Part 33). Walter de Gruyter, Berlin. 

Penny, N. D. 1977. Lista de Megaloptera, Neuroptera 
e Raphidioptera do Mexico, America Central, il- 
has Carafbas e America do Sul. Acta Amazonica 
7(4): Suplemento, 61 pp. 

. 1981 [1982]. Neuroptera of the Amazon Basin. 

Part 4, Sialidae. Acta Amazonica 1 1: 843-846. 

. 1982. Neuroptera of the Amazon Basin. Part 

7, Corydalidae. Acta Amazonica 12: 825-837. 

Penny, N. D., P A. Adams, and L. A. Stange. 1997. 
Species catalog of the Neuroptera, Megaloptera, 
and Raphidioptera of America north of Mexico. 
Proceedings of the California Academy of Scienc- 
es 50: 39-114. 

Penny, N. D. and O. S. Flint, Jr. 1982. A revision of 
the genus Chloronia (Neuroptera: Corydalidae). 
Smithsonian Contributions to Zoology 348: 1-27. 

Rambur, J. P. 1842. Histoire naturelle des insectes. 
Nevropteres. Librairie Encyclopedique de Roret, 
Paris. 

Ross. H. H. 1937. Studies of Nearctic aquatic insects, 
I. Nearctic alder flies of the genus Sialis. Illinois 
Natural History Survey Bulletin 21: 57-78. 

Stitz, H. 1914. Sialiden der Sammlung des Berliner 
Museums. Sitzungsberich der Gesellschaft natur- 
forschender Freunde zu Berlin 5: 191—205, 2 pis. 

Walker, F 1853. List of the specimens of neuropterous 
insects in the collection of the British Museum. 
Part II (Sialidae-Nemopterides). Edward Newman, 
London, pp. 193-476. 

. 1858. Characters of undescribed Neuroptera 

in the collection of W. W. Saunders. Transactions 
of the Royal Entomological Society of London, 
new series 5: 176-199. 

Weele, H. W., van der. 1909. New genera and species 
of Megaloptera Latr. Notes from the Leyden Mu- 
seum 30: 249-253. 

. 1910. Megaloptera (Latreille), monographic 

revision, pp. 1-93 + 4 pis. //; Collections Zoolo- 
giques du Baron Edm. de Selys Longchamps Ease. 
V (Premiere partie), Bruxelles. 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999, pp. 285-289 

IDENTITY OF SYRISTA SPECIOSA MOCSARY AND NOTES ON THE GENUS 
UROSYRISTA MA A (HYMENOPTERA: CEPHIDAE) 

David R. Smith 

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, MRC 168, Washington, DC 
20560-0168, U.S.A. (e-mail: dsmith@sel.barc.usda.gov) 



Abstract. — The holotype of Syrista speciosa Mocsary, described from Vietnam, is re- 
described and illustrated and confirmed as belonging to the genus Urosyrista Maa. Pos- 
sible characters are presented to separate it from other species of this southeastern Asian 
genus. 

Key Words: Cephidae, Urosyrista, stem sawflies 



In the treatments of world Cephidae 
(Benson 1946, Muche 1981) and Asian Ce- 
phidae (Maa 1944, 1949), the species de- 
scribed as Syrista speciosa by Mocsary 
(1904), an unusually large and colorful ce- 
phid from "Tonkin," has never been stud- 
ied and confidently placed. Benson (1946) 
placed S. speciosa in his new genus Ce- 
phalocephus, qualifying it with the state- 
ment that "it seems almost certain" that it 
belongs to this genus. Maa (1949) synon- 
ymized Cephalocephus with Urosyrista 
which he had described earlier (Maa 1944). 
Although Maa (1949) did not see Cephal- 
ocephus xanthiis Benson, the type species 
of Cephalocephus, both generic descrip- 
tions are almost identical and their synon- 
ymy cannot be disputed. Maa (1949) did 
not mention S. speciosa or put it in his key 
to species of Urosyrista. Muche (1981), 
who based much of his work on the liter- 
ature, placed S. speciosa in Urosyrista but 
did not include it in his key to Urosyrista 
species, which was taken directly from Maa 
(1949). 

Here, I redescribe, illustrate, and confirm 
the generic placement of Syrista speciosa 
based on examination of the holotype. 



Urosyrista speciosa (Mocsary) 
(Figs. 1-9) 

Syrista speciosa Mocsary 1904: 496. Fe- 
male. "Tonkin: Montes Mauson, in alti- 
tudine 2-3,000 pedum a H. Fruhstorfer 
detecta. (Mus. Hung.)." 

Cephalocephus speciosa: Benson 1946: 
100. 

Urosyrista speciosa: Muche 1981: 265. 

Description. — Length, 16 mm. Head and 
body smooth, shining, without punctures. 
Color: Yellowish with black markings (as 
shown in Figs. 1-9); antenna pale yellowish 
ventrally, black dorsally, with basal 3 seg- 
ments black. Wings hyaline, apices slightly 
blackish; most of stigma black, and veins 
dark brown, costa and margins of stigma 
yellowish. Head: In dorsal view, enlarged 
and swollen behind eyes, elongated, 1.2X 
broader than long, distance behind eyes 
1.3X eye length; distance from front ocellus 
to hind margin of head 3X distance from 
front ocellus to antennal sockets. Antennal- 
tentorial distance 2X distance between an- 
tennae. Antenna 30-segmented. Left man- 
dible bidentate, subapical tooth slightly lon- 
ger and stouter than apical tooth, without 



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Figs. 1-3. Urosyrista speciosa. \, Lateral view. 2, Dorsal view. 3, Dorsal view of thorax. 



VOLUME 101. NUMBER 2 



287 





■¥ 



f 



\ 




m 




Figs. 4-6. Urosyrista speciosa. 4. Head, front view. 5, Head, lateral view. 6. Head, dorsal view. 



288 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 











Figs. 7-9. Urosyrista speciosa. 7, Thorax, lateral view. 8, Apex of abdomen and siicaili. laicral view. 9, 
Apex of abdomen and sheath, dorsal view. 



intermediary tooth (Fig. 4). Sixth segment 
of maxillary palpus originating from near 
apex of 5th segment; segment 4 about 1.5X 
length of segment 6. Thorax: Forewing 
with anal crossvein. Hindwing with cubital 



cell. Hindtibia with 2 preapical spines. Mid- 
tibia with one preapical spine. Hindbasitar- 
sus subequal in length to remaining tarsal 
segments combined. Tarsal claws bifid, in- 
ner tooth about as long as outer tooth, with- 



VOLUME 101. NUMBER 2 



289 



out basal lobe. Abdomen: Cercus less than 
half length of sawsheath (Figs. 8, 9). Sheath 
as in Figs. 8, 9; in dorsal view, thick and 
parallel sided; length about .6X length of 
basal plate and slightly less than half as 
long as hind basitarsus. 

Holotype. — Female, labeled "Tonkin, 
Montes Manson, April, Mai, 2-3,000', H. 
Fruhstorfer," "Typus 1904 Syrista speciosa 
Mocs." (red label). The third label on the 
pin is a blank red label. In the Hungarian 
Natural History Museum, Budapest. 

Discussion. — Almost all characters of 
Syrista speciosa are consistent with Uro- 
syrista, and placement in this genus is cor- 
rect according to generic definitions and 
keys by Maa (1944, 1949), Benson (1946), 
and Muche (1981). Some minor differences 
are the lack of a basal swelling on the tarsal 
claws (according to Benson 1946 the claws 
have a small basal swelling) and the sub- 
apical tooth of the left mandible which, al- 
though longer than the apical tooth, is not 
as large in relation to the apical tooth as 
figured by Benson (1946, fig. 13). Unique 
characters for Urosyrista in the Cephidae 
are the enlarged head behind the eyes (Figs. 
5, 6), the lack of an intermediate tooth and 
large innter tooth of the left mandible (Fig. 
4), the apical segment of the maxillary pal- 
pus emerging from near the apex of the fifth 
segment, and the bifid tarsal claws, lacking 
a basal lobe. 

Urosyrista is known only from south- 
eastern Asia, and three species are currently 
included: U. speciosa (Mocsary) from Vi- 
etnam; U. montana (Maa) {— Cephaloce- 
phiis xanthus Benson according to Maa 
1949) from China and Burma; and U. men- 
cioyana Maa from China. Maa ( 1 949) con- 
sidered three color forms of U. mencioyana, 
the typical form, var. itnicolor Maa, and var. 
xanthobalteata Maa. All three are separated 
by the amount of black on the pronotum. 
The host plant is known only for U. men- 
cioyana; specimens were reared irom. Acan- 
thopanax trifoliatus (Lour.) Merr. (Arali- 
aceae) (Maa 1944). 

Urosyrista speciosa appears to be most 



similar to U. montana because they share 
the thick, parallel sided sheath in dorsal 
view. The sheath of U. mencioyana is slen- 
der and gradually tapering toward its apex. 
I hesitate to use coloration. In species as 
these with numerous black markings, vari- 
ation in the amount of black can be exten- 
sive. Maa ( 1949) has alluded to this by sep- 
arating several color forms of U. mencioy- 
ana. However, as it may be useful, U. spe- 
ciosa is more extensively yellow than the 
other two species and the antennae are 
black dorsally, pale whitish ventrally, with 
the basal three segment entirely black. Uro- 
syrista montana has the antennae apically 
dull brown to black, basally distinctly paler, 
and the mesepisternum mostly black with a 
median yellow band. Urosyrista mencioy- 
ana has the antennae apically yellow, ba- 
sally distinctly darker, and the head and tho- 
rax mostly black. 

Acknowledgments 

I thank L. Zombori, Hungarian Museum 
of Natural History, Budapest, for loan of 
the holotype. Terry Nuhn and Cathy An- 
derson, Museum Technicians, Systematic 
Entomology Laboratory, USDA, took the 
photographs of the holotype and arranged 
the plates. I thank the following for review 
of the manuscript: N. M. Schiff, USDA, 
Forest Service, Stoneville, MS, and M.E. 
Schauff and S.W. Lingafelter, Systematic 
Entomology Laboratory, USDA, Washing- 
ton, DC. 

Literature Cited 

Benson, R. B. 1946. Classification of the Cephidae 
(Hymenoptera Symphyta). Transactions of the 
Royal Entomological Society of London 96: 89- 
108. 

Maa, T. C. 1944. Novelties of Chinese Hymenoptera, 
Chalastogastra. Biological Bulletin of the Fukien 
Christian University 4: 33-60. 

Maa, T. C. 1949. A synopsis of Chinese sawflies of 
the superfamily Cephoidea (Hymenoptera). Chi- 
nese Journal of Zoology 3: 17-42. 

Mocsary, A. 1904. Siricidarum species quinque novae. 
Annates Musei Nationalis Hungarici 2: 496-498. 

Muche, H. 1981. Die Cephidae der Erde (Hym., Ce- 
phidae). Deutsche Entomologische Zeitschrift, 
N.E 28: 239-295. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999. pp. 290-294 

FRANKLINIELLA ZUCCHINI (THYSANOPTERA: THRIPIDAE), A NEW 
SPECIES AND VECTOR OF TOSPOVIRUS IN BRAZIL 

SuEO Nakahara and Rhnata C. Monteiro 

(SN) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, 
10300 Baltimore Avenue, Beltsville, Maryland 20705-2350, U.S.A. (e-mail: snakahar@ 
sel.barc.usda.gov); (RCM) Departamento de Entomologia, ESALQ, Universidade de Sao 
Paulo, 13418-900 Piracicaba, Sao Paulo, Brazil 



Abstract. — Frankliniella zucchini, new species, is described. It is a vector of a tos- 
povirus that causes zucchini lethal chlorotic disease of Cucurbita pepo L. cv. Caserta in 
Sao Paulo State, Brazil. 



Key Words: 



Frankliniella 
ZLCV, Brazil 



zucchini n. sp., Thysanoptera, Thripidae, zucchini, vector. 



A serious disease of zucchini squash, Cu- 
curbita pepo L. cv. Caserta, currently 
known as zucchini lethal chlorotic disease 
(ZLC), was discovered during 1995 in ex- 
perimental fields at Piracicaba, Sao Paulo 
State, Brazil (Rezende et al. 1997). This 
disease is cau.sed by a species of Tospovi- 
rus, zucchini lethal chlorotic virus (ZLCV) 
(Pozzer et al. 1996). Tospoviruses can be 
transmitted only by thrips adults and larvae. 
The disease apparently was present sporad- 
ically in the state prior to 1991 when many 
zucchini plants were observed with symp- 
toms of ZLC in Campinas county. Since 
then, symptoms of ZLC were observed 
more frequently on zucchini squash and 
watermelon. Frankliniella zucchini, new 
species, described here was the predomi- 
nant thrips species collected from foliage 
and flowers of infected plants in Piracicaba. 
In preliminary transmission tests, this thrips 
was found to be a vector of ZLCV to zuc- 
chini seedlings (Rezende 1998, personal 
communication). 

Two polyphagous Frankliniella species 
are vectors of "tomato spotted wilt virus" 
(TSWV) and "tomato chlorotic spot virus" 



(TCSV) in Brazil (Wijkamp et al. 1995). 
Frankliniella schnitzel (Trybom) dark form 
is an efficient vector of TSWV and TCSV 
that damages tomato crops. Although the 
yellow form of this species was previously 
not considered to be a vector, it is an inef- 
ficient vector of TSWV according to Wijk- 
amp et al. (1995). The recently established 
F. occidentalis (Pergande) is a pest of var- 
ious agricultural crops and also a vector. 
Frankliniella zucchini is known to vector 
only ZLCV and is the fifth Frankliniella 
species to be implicated in the transmission 
of tospoviruses. The other two are F. fusca 
(Hinds) and F. intonsa (Trybom). 

For measurements and counts, the values 
for the holotype are given first and are fol- 
lowed by values for the paratypes in paren- 
theses. If parentheses are absent the values 
are either for the holotype or for all mea- 
sured specimens. 

Frankliniella zucchini 

Nakahara and Monteiro, 

new species 

(Figs. 1-6) 
Female (macropterous). — Body generally 
yellow; forewing pale yellow, legs paler 



VOLUME 101, NUMBER 2 



291 




Figs 1-6. FmnkUniella zucchini. Female. 1, Right antenna. 2, Head (a, ocellar seta III; b, POiv seta). 3, 
Pronotum (a, antcromarginal seta; b, anteroangular setae; c, outer posteroangular seta; d, inner posteroangular 
seta, e. posteromarginal seta II; f, submarginal seta). 4, Metanotum. 5. Abdominal tergite VIII. 6, Abdominal 
tergite IX (a, Blseta; b, 82 seta; c, 33 seta). Scale = 0.1 mm. 



292 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



yellow than body; body and forewing setae 
brown. Antennal segment I as pale as head; 
segment II pale yellow in proximal Vi, shad- 
ed light brown distally; segment III pale 
yellow in proximal % including pedicel, 
distally brown; segment IV pale yellow in 
proximal Vi, brown distally and in pedicel; 
segment V pale yellow in proximal Vs, dis- 
tally brown; segments VI to VIII brown. 

Antenna (Fig. 1): More than twice as 
long as head; pedicel of segment III with 
slight angulation, segments III and IV each 
with V-shaped forked sense cones, 22-27 jx 
long; segment III distad of subapical setae 
slightly converging to apex, about V4 length 
of segment, segment IV constricted into a 
neck in distal part; segment VI slightly ped- 
icellate at base, inner sense cone 32-35 |jl 
long, extending distally at least to apex of 
segment VII. 

Head (Fig. 2): Slightly shorter than pro- 
notum, about 1.5 times wider than long, 
cheeks rather straight, compound eyes al- 
most twice as long as occiput; interocellar 
area without sculpturing, caudad of com- 
pound eyes with transverse striae, those 
more posterior spaced farther apart. Com- 
pound eyes each with 4 pigmented facets in 
1-2,3,5 pattern (see Nakahara 1988). Di- 
ameter of fore ocellus 15-17 (x. Ocellar se- 
tae I short, about 12 |x long, just anterior of 
fore ocellus; ocellar setae II short, laterad 
of fore ocellus and just mesad of inner mar- 
gin of compound eye, about 12 jx long; 
ocellar setae III well developed (Fig. 2a), 
between anterior part of posterior ocelli, 
separated by about diameter of fore ocellus, 
45(37-48) |x long. Postocular (PO) setae 5 
pairs; POi seta absent; POii seta 12-15 |jl 
long, caudad of posterior ocellus; next lat- 
erad POiii seta 10-12 |x long; POiv seta 
longest (Fig. 2b), 32(24-33) |x long; POv 
and POvi setae laterad of and shorter than 
seta iv but longer than setae ii and iii. 
Mouthcone conical, longer than head, distal 
-/; less convergent than basal X,, extending 
to posterior margin of prosternum; mandi- 
ble 124-136 jx long. 

Pronotum (Fig. 3): Rectangular, broader 



than long, sculptured with irregularly 
spaced transverse striae and sparsely anas- 
tomosing. Median discal area with 3-5 se- 
tae 10-12 (X long; 2 irregular rows of discal 
setae in posterior V3, 7-9 setae in anterior 
row including a pair of longer submarginal 
setae (Fig. 3f), 20-24 |x, anterior of 
posteroangular setae; posterior row normal- 
ly with 4 setae; anteromarginal setae 37- 
42(42-48) |x long (Fig. 3a), 2 short setae, 
10-12 |x long, between anteromarginal se- 
tae; anteroangular setae 48(48-59) |x long 
(Fig. 3b); posteroangular seta inner pair 59 
(57-67) fx long (Fig. 3d), outer pair 57- 
59(59-64) |x long (Fig. 3c), about V2 as long 
as notum; posteromarginal seta II 32(32- 
40) |x long (Fig. 3e). Mesonotum: Subtra- 
pezoidal, anterior angulate area smooth 
with pair of campaniform sensilla, median 
and posterior parts transversely sculptured, 
laterally with striae oriented longitudinally; 
2 pairs of short setae on or near posterior 
margin, inner pair 1 7—20 |x long, outer pair 
slightly stouter, 20-24 |x long. Metanotum 
(Fig. 4): Reticulated with most reticles lon- 
ger than wide and oriented longitudinally, 
reticles in anterior medial area more polyg- 
onal and wider than other reticles; median 
setae 55-59(54-59) |x long, thicker than 
lateral setae 35-37(35-37) |x long; 2 cam- 
paniform sensilla in posterior V-i of notum. 

Forewing: Rather straight, apex pointed; 
fringe cilia wavy; costa with 20-24(19-20) 
cilia, 23-24(22-24) setae, setae at mid- 
length 40(37-42) |x long, shorter than width 
of forewing; forevein with 19-21(18-20) 
setae, hindvein with 16-17(14-15) setae; 
scale with 4 marginal and 1 discal setae. 

Abdomen: Tergites sculptured anteriorly, 
and laterally of median setae and campan- 
iform sensilla; median setae short, on VII 
17-20 |x long; short ctenidia on tergite IV; 
posterior margins of intermediate tergites 
with series of low, truncate lobes, with a 
few small teeth laterally; tergite VIII with 
posteromarginal comb complete with 
14(13-17) microtrichia, longest 17-20 |x 
long (Fig. 5); tergite IX with microtrichia 
on anastomosing striae, posterior pair of 



VOLUME 101, NUMBER 2 



293 



campaniform sensilla near Bl setae (Fig. 
6); tergite X subequal to IX, dorsal split 
almost complete on X. Ovipositor well de- 
veloped, 183(222-235) fx long. 

Measurements: Female holotype and 
(paratypes) in |jl. Body length from anterior 
of eye 1176(1161-1221), distended 
1423(1408-1568). Antenna: Total length 
270(265-280); length and width of segment 
I 24(24), 27(27-30); II 37(35-37), 24(27); 
III 50(48-50), 22(22); IV 45(42-48), 
20(20-21); V 37(37-40), 17(17(20); VI 
52(52-54), 18(17-20); VII 10(10-11), 
7(7); VIII 15( 15), 5(5-6). Head length from 
anterior of compound eye 100(96-106), 
width at compound eyes 151(148-156), 
width at cheeks 156(143-158); length of 
compound eye 62(62-69), width 45; length 
of occiput posterior of compound eye 
37(35-37). Pronotal length 124(126-133), 
width 190(178-190). Forewing length 
729(679-729), width at midlength 57(52- 
57). Length of abdominal tergite IX 62(64- 
67), length of Bl setae 104-109(96-109) 
(Fig. 6a), B2 setae 101-106 (104-109) 
(Fig. 6b), B3 seta 100(100) (Fig. 6c); length 
of tergite X 64(64-67), length of Bl seta 
109(96-1 1 1), B2 seta 98(94). 

Male (macropterous). — Smaller than fe- 
male, otherwise similar in color and most 
anatomical structures. Body length 1,000- 
1,050 |x. Antennal length 220-246 jx. Ab- 
dominal tergite VIII with complete postero- 
marginal comb with 13-14 long, slender 
microtrichia; sternites III- VII each with 
transversve glandular area with anterior and 
posterior margins concave, on III 52-62 |x 
wide, 15-17 |x long, on VII 40-54 |jl wide, 
12-15 |x long, 0.30-0.38 as wide as ster- 
nite; sternite VIII with posteromarginal mi- 
crotrichia. 

Type material. — Holotype 9 and 18 $ 
and 2 d paratypes: Brasil, Sao Paulo, Pir- 
acicaba, Cucurbita pepo L, cv. Caserta, 14- 
VII-95. R.C. Monteiro. Holotype and 10 
paratypes deposited in Departamento de 
Entomologia, ESALQ, Universidade de Sao 
Paulo, Piracicaba, Brazil, 10 paratypes in 
the National Museum of Natural History, 



Smithsonian Institution, Washington D.C, 
and 3 paratypes in The Natural History Mu- 
seum, London, United Kingdom. 

Etymology. — The species is named after 
the common name of the host, "zucchini," 
and is a noun in apposition. 

Distribution. — Known only from Sao 
Paulo State, Brazil. 

Collected from. — Cucurbita pepo L. cv. 
Caserta (zucchini). 

Comments. — Frankliniella gemina Bag- 
nail and F. rodeos Moulton in Brazil are 
similar to F. zucchini in color and most an- 
atomical characters. Frankliniella zucchini 
lacks POi seta and has ocellar setae III po- 
sitioned between the anterior part of pos- 
terior ocelli and separated by about the 
width of the fore ocellus; whereas POi setae 
are present in the other two species and 
ocellar setae III are aligned with the ante- 
rior margin of posterior ocelli or slightly 
anterior and are farther apart. 

Most Frankliniella species have three 
pairs of short postocular (PO) setae mesad 
or anteromesad of the fourth or longest pair 
of PO setae which is positioned posterior of 
the compound eye (Fig. 2). When only two 
pairs of short PO setae are present, the POi 
seta is normally absent. The normal posi- 
tion of POi seta is caudad of and usually 
slightly mesad of the inner margin of the 
posterior ocelli. 

Acknowledgments 

We thank the following colleagues for 
their contributions to this article: Laurence 
Mound, CSIRO, Canberra, Australia, rec- 
ognized that this thrips was new to science 
and recommended that it be described; and 
J. A. M. Rezende, ESALQ/USP, Piracicaba, 
Sao Paulo, Brazil, provided information 
about ZLCV and the vector For reviewing 
the manuscript and useful suggestions, we 
thank Robert Foottit, Agriculture and Agri- 
Food Canada, Ottawa, and Stuart Mc- 
Kamey and Douglass Miller, Systematic 
Entomology Laboratory, USDA, Washing- 
ton D.C. and Beltsville, Maryland respec- 
tively. 



294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

Literature Cited Resende, A. C. De Avila, and S. M. Scagliusi. 

., , , „ ,„oo n 1 ■■ J c r I • 1997. Incidence, biological and serological char- 

Nakahara, S. 1988. Preliminary study of racetal pig- 

, ,. • ,, , e T u .■ acteristics of a tospovirus infecting experimental 

mentation in the compound eyes of Terebrantia ^ t. h- 

(Thysanoptera). Acta Phytopathologica et Ento- ^^^^^ "^ ^"^'^h'"' '" ^^^ ^^"1° ^'"'^- ^'-azil- Fi- 

mologica Hungarica 23(3-4): 321-329. topatologia brasileira 22: 92-95. 

Pozzer, L., R. O. Resende, M. I. Bezerra, T Nagata, Wijkamp, I., N. Almarza and D. Peters. 1995. Median 

M. I. Lima, F W. Kitajima, and A. C. De Avila. latent period and transmission of tospoviruses 

1996. Zucchini lethal chlorotic virus (ZLCV), a vectored by thrips. pp. 153-156. //; Parker, B. L., 

proposed new species in Tospovirus genus. Fito- M. Skinner and T Lewis, eds., Thrips Biology and 

patologia Brasileira 21 (Suplemento): 432. Mangement. Plenum Press, New York and Lon- 

Rezende, J.A.M., S. R. Galleti, L. Pozzer, R. de O. don. 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999, pp. 295-299 

THE LARVA OF CHALCIDOMORPHINA AURATA ENDERLEIN 1914 

(DIPTERA: STRATIOMYIDAE) FROM "ILHA DE MARAMBAIA," 

RIO DE JANEIRO, BRAZIL 

JO.SH R. PUJOI.-LUZ AND ROBKR TO DI-; XliREZ 

Departamento de Biologia Animal, Instituto de Biologia da Universidade Federal do 
Rio de Janeiro, Seropedica, Rio de Janeiro, 23890-000, Brazil (e-mail: pujol-luz@uol. 
com.br; rdexerez@uol.com.br) 



Abstract. — The larva of Chalciclomorphina aurata is described for the first time, based 
on six larvae and the puparium. Larvae were collected under the bark of fallen trees in 
a tropical rain forest at Ilha da Marambaia, State of Rio de Janeiro, Brazil (23°04'S, 
43°53'W, approximately 42 km-). Some biological notes are also presented. 

Key Words: Stratiomyidae, Pachygastrinae, Chalciclomorphina aurata, soldier-flies, lar- 
vae, tropical rain forest 



The pachygastrine genus Chalciclomor- 
phina Enderlein, 1914, with four species, is 
widespread in the Neotropics, from Mexico 
to Brazil: Chalcidc^morphina aurata Ender- 
lein, 1914 (Mexico, Panama, Colombia, 
Peru and Brazil); C. plana James, 1967 
(Dominica); C terataspis James, 1974 
(Peru); and C. argentea McFadden, 1980 in 
James et al., 1980 (Mexico) (James 1973, 
James et al. 1980). 

James (1974) recognized Chcdciclcjmor- 
phina based on the following characters: ( 1 ) 
antenna elongate, with long scape and fla- 
gellum, and (2) scutellum projected into an 
elongate, spur-like process (Fig. 1 ). James 
( 1 974) segregated Chalcidomorphina and 
Dactylaccmtha Lindner, 1964, from the re- 
lated genus Dactylodeictes Kertesz, 1914, 
based on wing venation. 

Pachygastrinae larvae from the Neotrop- 
ics have never been described. McFadden 
(1967) and James (1981) furnished the last 
records of the known North American lar- 
vae of this subfamily. They studied Nearc- 
tic representatives of some genera that are 



widespread over much of the Americas 
(e.g., Eididimits Kertesz, 1914 [ = Eucyni- 
pimorpha Malloch, 1915; =Eiipachygaster 
authors, part, not Kertesz, 1911]; Gowdey- 
ana Curran, 1928 [^Eiipachygaster au- 
thors, part, not Kertesz, 191 1; =Paraeidal- 
imus Lindner, 1964] and Zahrachia Co- 
quillett, 1901) (James et al. 1980). 

Here we describe the larva of Chalcido- 
morphina aurata for the first time, based on 
six larvae and the puparium. Some of the 
larval features employed in this work are 
the same used by McFadden (1967) and 
James (1981) to describe other genera. 

Chcdcidomorphina larvae were collected 
under the bark of fallen trees in a tropical 
rain forest at Ilha da Marambaia, State of 
Rio de Janeiro, Brazil, (23°04'S, 43°53"W, 
approximately 42 km-) outside and inside 
the forest behind a dam (a restricted area of 
the Brazilian Navy). 

The junior author collected approximate- 
ly 35 to 50 larvae of different instars at 
each site belonging to two genera, Chcdci- 
domorphina and Cyphomyia Wiedemann, 
1819 (Clitellariinae). Field and laboratory 



296 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





Figs. 1-2. Chalcicioniorphinci auntlci. 1, Female habitus (all pilosity is omitted). 2. Larva (right in dorsal 
view; left in ventral view). Scale line = 2.0 mm. 



observations suggest that larvae of C. aii- 
rata feed on microorganisms occurring in 
the moist areas beneath the bark of trees. 
The trees where the larvae were found were 
not identified. We also confirm McFadden's 
(1967) statement concerning the gregarious 
behavior of larvae of pachygastrine soldier- 
flies. 

The larvae of C. ourata were reared in 
semi-natural conditions, segregated by size 
classes, and put on individual petri dishes 
having as substrate the same moist sub- 
stance present where the larvae were living. 
A single female was collected flying over a 



tree. We believe that oviposition occurs in 
the moist substrate, through crevices in the 
bark, because a large number of the small- 
est larvae of C. aurata were found there. 

The females that emerged are extremely 
similar to those described by Lindner 
(1951), James (1974) and James et al. 
(1980), but showed a chromatic pattern of 
the eyes not described by these authors. The 
eyes in living insects are brownish with a 
greenish "9-shaped" pattern in lateral view 
(Fig. 1). 

The terminology adopted in the descrip- 
tions follows James (1981) and Rozkosny 



VOLUME 101, NUMBER 2 



297 






Figs. 3-9. Larva of Chalcicloniorphiini aiinifa. 3, Head, dorsal view. 4, Head, ventral view. 5, Right spiracle, 
lateral view. 6, Abdominal segment 6, dorsal view. 7, Abdominal segment 6, ventral view. 8, Abdominal segment 
8, dorsal view. 9, Abdominal segment 8, ventral view. Scale lines = 0.5 mm (Figs. 3-5), = 1.0 mm (Figs. 6- 
9). Abbreviations: ant = antenna; cyb = cylindrical brushes; Ibr = labrum. 



298 



PROCHliDINGS Ol' THli ENTOMOLOGICAL SOCIETY OF WASHINGTON 



and Kovac (1994). The specimens upon 
which this study is based are in the Colevao 
Entomologica Costa Lima (CECL), Insti- 
tute) de Biologia, Universidade Federal Ru- 
ral do Rio de Janeiro (UFRRJ), Seropedica, 
Rio de Janeiro, RJ, Brasil. 

Chalcidomorphina aunifa Enderlein, 1914 
(Figs. 1-9) 

Chalcidomorphina aurata: Enderlein 1914 
(original description); Lindner 1951 
(suppl. descr., figs.); James 1973 (cata- 
logue); James 1974 (revision, figs.); 
James, McFadden, and Woodley 1980 
(suppl. descr., key to females, notes on 
the males, figs.). 

Distribution. — Neotropical: Mexico, 
Panama, Colombia, Peru, Brazil. 

Larva (and puparium). — Length 5.0 to 
5.3 mm, flattened dorsoventrally, lateral 
margins of body segments strongly arched. 
Cuticle with usual mosaic appearance, 
some cells forming characteristic patches 
and plaques on abdominal segments 6 and 
8. Chromatic pattern yellowish brown, with 
some dark punctuations (Fig. 2). 

Head: Subconical, moderately flat; 
mandibular-maxillary complex with well 
developed, cylindrical brushes almost as 
long as labrum, in dorsal view (Fig. 3); la- 
brum triangular Antenna short, rising at an- 
terior part of head. Eyes prominent, round- 
ed, arising at the posterior part of the head. 
Two pairs of lateral setae, one pair of cly- 
peofrontal setae and one pair of dorsolateral 
setae inserted above eyes; three pairs of 
ventrolateral setae and three pairs of spine- 
like ventral setae (Figs. 3-4). 

Thorax: First segment shorter than oth- 
ers. Spiracle prominent and V-shaped, with 
a small anterolateral spiracular seta, in lat- 
eral view with two dorsal setae and one 
ventral seta (Fig. 5). First segment with two 
rows of setae in dorsal view: two pairs of 
anterodorsal setae and three pairs of dorsal 
setae. In ventral view, two pairs of ventral 
setae and one pair of ventrolateral setae 
near spiracle. Second and third segments 



with one row of setae with four pairs of 
dorsal setae and three pairs of ventral setae 
(Fig. 2). 

Abdomen: Segments 1-7 similar in 
shape to thoracic segments (Fig. 2), with a 
row of five pairs of dorsal setae and four 
pair of ventral setae (Fig. 6-7); ventrome- 
dial line of segment 6 with an elliptical ster- 
nal patch (Fig. 7); segment 8 rounded, with 
three pairs of conspicuous plaques along 
dorsomedial line between a pair of pennate 
dorsocentral setae (Fig. 8), four pairs of lat- 
eral setae, apical pair shorter than others; 
opening of spiracular chamber with a fringe 
of small setae, anal slit on ventral side 
emarginate with a long fringe of setae on 
each side (Fig. 9). 

Material examined. — Brazil, Rio de Ja- 
neiro, Ilha da Marambaia, 17.n.l998 (R. 
Xerez col.), 4 females (emerged: 04.III.1998; 
2.IV.1998 and 9.1 V. 1998) and 2 larvae (last 
instar). 

Comments. — Chalcidomorphina larvae 
share some features with the genera keyed 
by James ( 198 1 ). The larva keys to the sec- 
ond half of couplet 21. Then, the features 
are distributed in several couplets. Couplet 
21 segregates two groups: first half: [Berk- 
shiria + Neopachygaster] and second half: 
[Gowdeyana + Eidalimus + Pachygaster 
+ Zahrachia]. However, Chalcidomorphina 
also shares a feature with Neopachygaster 
(second half of couplet 22, three pairs of 
conspicuous plaques along dorsomedial line 
of abdominal segment 8) and differs in the 
same couplet by the shape of sternal patch 
of abdominal segment 6 (oval in Neopach- 
ygaster). 

Ac'KNOWLIiDC.MLNTS 

We thank to Dr. Nelson Papavero, Dr. 
Francisco Racca Filho, Dr. Helio Ricardo 
da Silva, Mary S. Linn, and Christopher 
Shell for critical reviews of the manuscript. 
This research was financed by the Conselho 
Nacional de Desenvolvimento Cientifico e 
Tecnologico— CNPq (Grant 300265/96-4). 



VOLUME 101. NUMBER 2 



299 



LiTERAIURK CiTHD 

James, M. T. 1973. A catalogue of the l)ip(era of the 
Americas south of the United States. 26. Family 
Stratiomyidae. Museu dc Zoologia da Univeisi- 
dade de Sao Paulo. 95 pp. 

. 1974. The pachygastrine genera Dcu tylodeic- 

tcs, Chalciiloinorphina, and Thopomyict in South 
America (Diptera, Stratiomyidae). Melanderia 14: 
23-.'^2. 

. 1965. Stratiomyidae, pp. 299-.^ 19. In Stone, 

A., C. W. Sabrosky. W. W. Wirth, R. H. Poole, 
and J. R. CouLson, eds. A Catalog of Diptera of 
America North of Mexico. United States Depart- 
ment of Agriculture, Agriculture Handbook No. 
276. Washington, D.C. iv + 1696 pp. 

. 1981. Stratiomyidae. pp. 497-511. //; Mc 



Alpine, J. E, B. V. Peterson, G. E. Shewell, H. J. 
Teskey, .1. R. Vockeroth, and D. M. Wood, eds. 
Manual of Nearctic Diptera. Vol. 1. Agriculture 
Canada, Ottawa, vi + 674 pp. 

James, M. T, M. W. McFadden, and N. E. Woodley. 
1980. The Pachygasirinae (Diptera, Stratiomyi- 
dae) of Middle America. Melanderia 34: 1-36. 

Lindner, E. 195 1 . Vierter Beitrag zur Kenntnis der sue- 
damerikani.schen Strationiyiidenfauna (Dipt.). Re- 
vista de Entomologia 22 (1-3). 

McFadden, M. W. 1967. Soldier Fly larvae in America 
North of Mexico. Proceedings of the United States 
National Museum 121 (3569): 1-72. 

Ro/.kosny, R. and D. Kovac. 1994. Adults and larvae 
of two Ptccliciis Loew from Peninsular Malaysia 
(Diptera, Stratiomyidae). Tijdschrift voor Enlo- 
mologie 137: 75-86. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999, pp. 300-311 

DESCRIPTION OF THE IMMATURE STAGES OF THREE SPECIES OF 

EULEPIDOTIS GUENEE (LEPIDOPTERA: NOCTUIDAE) WITH 

NOTES ON THEIR NATURAL HISTORY 

Michael G. Pogue and Annette Aiello 

(MGP) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History, Washington, DC 
20560-0168, U.S.A. (e-mail: mpogue@sel.barc.usda.gov); (AA) Smithsonian Tropical Re- 
search Institute, P.O. Box 2072, Balboa, Ancon, Republic of Panama (e-mail: 
aielloa@tivoli.si.edu) 



Abstract. — Larvae and pupae of the genus Eiilepidotis Hiibner are described for the 
first time. The species are E. merricki (Holland), E. juncida (Guenee), and E. superior 
(Guenee), all of Neotropical distribution. The larval host of E. merricki is Spanish-lime, 
Melicocciis bijiigatus Jacq. (Sapindaceae), which is cultivated as an ornamental and fruit 
tree throughout the Caribbean. Larvae of E. juncida were reared from Inga fagifolia (L.) 
Willd. ex Benth. (= Inga laurina (Sw.) Willd.) (Mimosaceae). Larvae off. superior were 
defoliating Quararibea asterolepis Pitt. (Bombacaceae). 

Key Words: Melicoccus, Sapindaceae, Inga, Mimosaceae, Quararibea, Bombacaceae, 
host plants. Neotropical, Panama, Puerto Rico 



Larvae of tropical Lepidoptera are poorly 
known, and unless reared to adults, they are 
virtually impossible to identify specifically. 
We describe for the first time the immature 
stages of Eulepidotis Hiibner, a large genus 
belonging to the subfamily Catocalinae, 
that is mainly of Neotropical affinity and 
which comprises 105 species, 3 of which 
occur in North America (Poole 1989, Poole 
and Gentili 1996). The immature stages of 
E. merricki (Holland), E. juncida (Guenee), 
and E. superior (Guenee) were studied. 

There have been two reports of Eulepi- 
dotis larvae defoliating tropical trees. Wong 
et al. (1990) reported larvae of E. superior 
defoliating Quararibea asterolepis Pitt. 
(Bombacaceae) on Barro Colorado Island, 
Panama. Nascimento and Proctor (1994) re- 
ported that larvae of E. phrygionia Hamp- 
son were defoliating a monodominant rain- 
forest of Peltogyne gracilipes Ducke (Cae- 



salpiniaceae) on Maraca Island, Roraima, 
Brazil. Unfortunately no larvae from the 
latter study were preserved. 

Larvae, pupae, and adults of E. merricki 
were sent to one of us (MGP) by Lionel- 
Pagan, U.S.D.A., Animal and Plant Health 
Inspection Service, Plant Protection and 
Quarantine, San Juan, Puerto Rico, for 
identification. The larvae were defoliating 
Spanish-lime, Melicoccus bijugatus Jacq. 
(Sapindaceae), a tree used for fruit and as 
ornamental purposes in the Caribbean. 
Spanish-lime is native to continental tropi- 
cal America from Nicaragua to Surinam 
and is planted widely and becoming natu- 
ralized in the Caribbean (Adams 1972, 
Proctor 1984). 

One of us (AA) reared the immature 
stages of E. juncida and E. superior. The 
host of E. juncida was Inga fagifolia (L.) 
Willd. ex Benth. (= Ingalaurina (Sw.) 



VOLUME 101, NUMBER 2 



301 









**^*^^*m-F*i 



,v.^. 







Figs. 1-3. Larval habitus. 1, Eiilepidotis merricki. 2. E. juncida. 3, E. superior. 



Willd.) (Mimosaceae). The host of Eulepi- 
dotis superior is Quararibea asterolepis 
Pitt (Bombacaceae). 

Methods and Materials 

Twenty-two larvae of E. juncida were 
collected and designated as Aiello Lot 80- 
003. Fifteen of the larvae were preserved 
in 80% ethanol. Seven were placed in a cy- 
lindrical rearing cage along with both old 
and young foliage. The cage was construct- 
ed from petri dishes and aluminum window 
screen with a circle of paper towel on the 
floor and measured 10 cm tall by 9 cm in 
diameter. To maintain and regulate humid- 
ity, a damp, folded strip of paper towel was 
placed on the cage cover, and the entire as- 
sembly was kept inside a clear plastic 
ZipLoc® bag. After pupation the pupae 
were placed into separately numbered cages 
to await eclosion. Fifteen larvae, one pupa, 
and four adults with associated pupal skins 
are in the collection of the National Muse- 
um of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 



Three larvae and 10 pupae off. superior 
were obtained and designated as Aiello Lot 
85-19. The larvae were preserved in 80% 
ethanol, and seven adults with associated 
pupal skins were reared and are in the col- 
lection of the Smithsonian Tropical Re- 
search Institute, Republic of Panama. 

Key to Known Eulepidotis Larvae 

1 . Head dark brown to black, reticulate pattern 
reduced; mid-dorsal stripe black, solid (Fig. 1 ); 
setae A I — A2 — A3 form an acute angle (Fig. 

5) E. merricki (Holland) 

- Head brown, reticulate pattern covers most of 
head; mid-dorsal stripe broken (Figs. 2-3); se- 
tae Al — A2 — A3 form an obtuse angle (Figs. 
12-19) 2 

2. Head with reticulate pattern restricted to lateral 
area; dorsolateral markings on abdomen in 
pairs (Fig. 2); labrum with one pair of setae on 
lateral margin (Fig. 14) .... E. juncida (Guenee) 

- Head uniformly covered with a reticulate pat- 
tern; abdominal segments with single large 
dorsal markings (Fig. 3); labrum with two pairs 
of setae on lateral margin (Fig. 21) 

E. superior (Guenee) 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Key TO Known Eulepidotis Pupae 

1. Length less than 12 mm; labial palpus disjunct 
(Fig. 30) E. jitncida (Guenee) 

- Length greater than 13 mm; labial palpus con- 
tinuous (Figs. 25 and 34) 2 

2. Profemur absent (Fig. 25) 

E. merricki (Holland) 

- Profemur present (Fig. 34) 

E. superior (Guenee) 

Eulepidotis merricki (Holland) 

(Figs. 1, 4-10, 25-29) 

Larva 

Diagnosis. — Dorsal black stripe from 
pronotum to transverse stripe on segment 8 
and extending beyond band to segment 9. 
Head black to dark brown, without reticu- 
late pattern. 

Description. — Head (Figs. 5-10): Width 
of head capsule 1.8 ± .06 mm (range, 1.8- 
1.9 mm) (n = 5). Black to dark brown; ec- 
dysial line, epicranial suture, and posterior 
margin of head to stemmatal area cream. 
Prothoracic shield dark brown; medial 
patch dark brown withthin cream stripe; 
dorsolateral band cream; lateral edges dark 
brown. Labrum medially cleft; 3 pairs of 
dorsal setae forming an oblique line medi- 
ally (Fig. 7); 3 pairs of ventral epipharyn- 
geal setae (Fig. 8). Mandible with 3 distinct 
broad teeth; oral surface with broad molar- 
bearing process (Fig. 10). 

Thorax: Dorsal stripe black; dorsolateral 
band cream; lateral band broad, black, di- 
vided by a series of cream spots, extending 
to just below setae LI and L3. Legs dark 
brown. Underside dark brown to metatho- 
racic legs, caudal half of mesothorax cream; 
VI setae surrounded by dark brown spot. 

Abdomen (Fig. 1): Dorsal stripe black to 
broad transverse band on segment 8 and ex- 
tends between segments 8 and 9; dorsolat- 
eral band cream; lateral stripe black; lower 
lateral stripe with irregular margins, cream; 
spiracular stripe black with irregular mar- 
gins and not enclosing spiracle; below spi- 
racles and venter cream. Prolegs cream; 
plantae black; crochets in an uniordinal me- 
soseries. Segments 1 and 2 with 3 SV setae. 



Pupa 

Diagnosis. — Labial palpi present, contin- 
uous; profemur absent. 

Description. — Male (Figs. 25-27).- 
Length 15.8 ± 1.04 mm (range, 15.0-17.0 
mm) (n = 3). Labial palpi present, contin- 
uous. Profemur absent. Mesothoracic leg- 
reaching eye. Wings do not extend beyond 
caudal margin of segment 4. Segments 1-3 
and 8 with shallow circular pits on dorsum; 
segments 4-7 with shallow circular pits in 
a dense band extending about Va width of 
segment caudally and completely encircling 
segments 5-7; segment 9 smooth. Genital 
opening on a circular plate. Anal opening 
below genital opening. Cremaster consist- 
ing of a large median pair of slightly curved 
hooks and 3 pairs of small hooks, 1 pair 
located mediodorsally, 2 pairs located lat- 
erally (Fig. 27). 

Female (Figs. 28-29): Similar to male 
except: length 15.2 ± 0.35 mm (range, 
15.0-15.5 mm) (n = 2). Genital opening at 
caudal border of segment 8, dividing seg- 
ment 8 ventrally. Anal opening well caudad 
of genital opening. 

Host. — Melicoccus bijugatus Jacq.; Fam- 
ily Sapindaceae; common names: Spanish- 
lime, genip, honeyberry, mamoncillo, que- 
nette, Quenepa, Hongibeere. 

Natural history. — The only information 
known is that thousands of larvae were de- 
foliating the host plant (Lionel Pagan, per- 
sonal communication). 

Discussion. — The adult of E. merricki 
most closely resembles Eulepidotis carcis- 
tola Hampson and E. fumata (Felder and 
Rogenhofer). In E. merricki the prothorax 
and tegula have a black stripe; this is absent 
in E. carcistola and E. fumata. An elongate 
white spot at the base of median line of the 
forewing is larger and more distinct in E. 
carcistola than in E. merricki and E. fu- 
mata. The forewing lines in E. fumata are 
very faint compared with the bold distinct 
lines in E. merricki. 

Eulepidotis merricki was described from 
a specimen collected flying around low 



VOLUME 101. NUMBER 2 



303 




Figs. 4-10. Eidepidotis merricki larva. 4, Setal map. 5, Head. 6, Head, lateral view (scale = 0.5 mm). 7, 
Labrum, dorsal view. 8, Labrum, ventral view. 9, Mandible, dorsal view. 10, Mandible, ventral view (scale = 
0.1 mm). 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




11 



T2-3 



A 1-2 



A 3-6 




12 





15 




17 ^^ 



Figs. 11-17. Eiilepiilotis jimciila larva. 11, Setal map. 12, Head. 13, Head, lateral view (scale = 0.5 mm). 
14, Labrum, dorsal view. 15. Labriim. ventral view. 16, Mandible, dorsal view. 17, Mandible, ventral view (scale 
= 0.1 mm). 



herbage outside New Brighton, Pennsylva- onym of Palindia fumata Felder and Ro- 

nia, on August 5, 1900. Holland (1902) in genhofer (= E. fumata). The point is that 

the original description stated that it is very E. merricki is not a Nearctic species but a 

close io Palindia mabis GxxenQQ {= E. jun- Neotropical one, and that H. D. Merrick 

cida (Guenee)) which he confused as a syn- just happened to collect a specimen that 



VOLUME 101, NUMBER 2 



305 



was somehow imported from the Neotrop- 
ics, possibly on tropical fruits. There are no 
other specimens of E. merricki from the 
Nearctic in the National Museum, Washing- 
ton, D.C., or The Natural History Museum, 
London, collections. In Franclemont and 
Todd (1983) E. merricki is noted as being 
of questionable occurrence in America 
north of Mexico. 

Eiilepidotis merricki has a Caribbean dis- 
tribution, having been collected from Ja- 
maica, Cuba, and Puerto Rico. 

Eulepidotis jimcida (Guenee) 

(Figs. 2, 11-17, 30-33) 

Larva 

Diagnosis. — Head brown with reticulate 
pattern restricted to lateral area. Dorsum of 
abdomen with pair of dorsolateral brown 
patches encompassing setae Dl and D2. 

Description. — Head (Figs. 12-17): 
Width of head capsule for three instars as 
follows: 1.2 mm (n = 1); 1.8 ± .03 mm 
(range, 1.7-1.8 mm) (n = 7); 2.0 ± .05 mm 
(range, 2.0-2.1 mm) (n = 7). Brown with 
lateral reticulate pattern; adfrontal area 
from ecdysial line to just beyond setae AFl 
cream; setae PI and P2 with cream pina- 
cula. Labrum medially cleft; 3 pairs of dor- 
sal setae with second pair below other pairs; 
3 pairs of ventral epipharyngeal setae (Fig. 
15). Mandible with 2 distinct broad teeth, 1 
smaller tooth mostly obscured from dorsal 
view; oral surface with a broad molar-bear- 
ing process with small pointed processes on 
either side (Fig. 17). 

Thorax: Prothoracic shield brown; dorso- 
lateral band cream; small cream patch be- 
tween D2 and XD2. Dorsal stripe or patch 
on segment 2 brown, on segment 3 cream; 
dorsolateral band cream; lateral band broad, 
brown, interrupted by several cream spots, 
extending to just below setae LI and L3. 
Legs brown. Underside mostly cream, 
brown shading between coxae and encom- 
passing VI setae. 

Abdomen (Fig. 2): Dorsum with pair of 
dorsolateral brown patches encompassing 
setae Dl and D2; lower lateral stripe cream, 



consisting of irregular spots and stripes that 
are more or less contiguous to segment 6, 
line not contiguous to absent on segments 
7 to 9; spiracular band brown, broad, ex- 
tending below LI seta; below LI seta and 
venter cream. Prolegs cream to brown lat- 
erally; plantae cream; crochets in an unior- 
dinal mesoseries. Segments 1 and 2 with 3 
SV setae. 

Pupa 

Diagnosis. — Small, less than 12 mm 
long. Labial palpi present, but disjunct. Pro- 
femur present. 

Description. — Male (Figs. 30-33): 
Length 9.3 ± 0.40 (range, 8.9-9.7 mm) (n 
= 3). Labial palpi present, disjunct. Profe- 
mur present. Mesothoracic leg reaching 
eye. Wings do not extend beyond caudal 
margin of segment 4. Segments 1-3 and 8 
dorsum with shallow circular pits; segments 
4-7 with shallow circular pits in a dense 
band extending about Va width of segment 
caudally and completely encircling seg- 
ments 5-7; segment 9 smooth. Genital 
opening on a circular plate. Anal opening 
caudad of genital opening. Cremaster con- 
sisting of a large median pair of curved 
hooks and 3 pairs of small hooks, 1 pair 
located mediodorsally, 2 pairs located lat- 
erally (Figs. 32-33). 

Female (Fig. 33): Similar to male except: 
length 9.4 ± 0.99 mm (range, 15.0-17.0 
mm) (n = 2). Genital opening at caudal 
border of segment 8, dividing segment 8 
ventrally. Anal opening well caudad of gen- 
ital opening. 

Host. — Inga fagifolia (L.) Willd. ex 
Benth. (= Inga laurina (Sw.) Willd.); fam- 
ily Mimosaceae. 

Natural history. — On March 5, 1980, an 
outbreak of larvae occuned on Barro Col- 
orado Island, Republic of Panama, near the 
900 meter mark on Thomas Barbour Trail 
on the central plateau of the island. Larvae 
were so abundant that the rainlike sound of 
their fecula landing on the leaf litter could 
be heard many meters away. 

Although the majority were in the cano- 



306 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



SDl\^ 


-^ 


.-^°' 


y^ -> 


^"^—801 


LI.-, 
SV1. 

V1» 


iy 


\ PP1 


-^vV 


C^" 



A10 




Figs. 18-24. Eulep'ulotis superior larva. 18, Setal map. 19, Head. 20, Head, lateral view (scale = 0.5 mm). 
21, Labrum, dorsal view. 22, Labnim, ventral view. 23, Mandible, dorsal view. 24, Mandible, ventral view (scale 
= 0.1 mm). 



VOLUME 101, NUMBER 2 



307 



py, larvae fed at various levels in the tree 
eating only young leaves. They dropped on 
silk lines to the forest floor and surrounding 
vegetation to molt. Molting took place un- 
der a sheet of silk across a leaf. Larvae 
walked by "looping," as do geometrids, but 
they have the full complement of five pairs 
of prolegs. 

Some larvae dropped, presumably in re- 
sponse to some perceived danger, and be- 
gan reascending almost immediately. They 
progressed slowly by gathering the silk line 
into a ball using their thoracic legs. Upon 
completion of their journey, they aban- 
doned the ball of silk. 

Seven larvae were brought into the lab- 
oratory on March 5; one died on March 7. 
The remaining six molted on March 9, and 
late on March 10 began preparing pupation 
chambers of fecula and intact leaves on the 
cage floor. Five pupated on March 1 1 , and 
the sixth larva failed to pupate and died on 
March 12. Two adults eclosed during the 
night of March 19-20, and two more 
eclosed during the night of March 20-21. 
The fifth pupa died and was preserved. Pu- 
pal duration was 9 days for females and 10 
days for males. 

Discussion. — In the adult stage, E. jiinci- 
da is similar to E. juliata (Stoll). In E. jim- 
cida the median brownish-yellow stripe ex- 
tends from the costa to the inner margin of 
forewing, in contrast to E. jimcida, which 
has a more whitish-yellow median stripe that 
does not extend to the inner margin. 

Eulepidotis jimcida has a wide distribu- 
tion from Mexico to Panama in Central 
America and from Guyana, Venezuela, and 
Colombia to Bolivia in South America. Eu- 
lepidotis juliata is known, so far, only from 
Guyana, Venezuela, and Bolivia. 

Eulepidotis superior (Guenee) 

(Figs. 3, 18-24, 34-37) 

Larva 

Diagnosis. — Reticulated pattern covering 
entire head. A single large mid-dorsal light 
brown (in alcohol specimens) mark on all 
abdominal segments. 



Description.— //eo^ (Figs. 19-24): 
Width of head capsule for two instars as 
follows: 1.9 mm (n = 1); 2.6 mm (n = 2). 
Brown with cream reticulated pattern. La- 
brum cream, medially cleft; 3 pairs of dor- 
sal setae forming a straight line medially; 2 
pairs of small setae along lateral edge; 3 
pairs of ventral epipharyngeal setae (Fig. 
22). Mandible with 2 outer setae; cutting 
surface with 3 indistinct teeth; oral surface 
with broad molar-bearing process (Fig. 24). 

Thorax: Prothoracic shield brown; mid- 
dorsal stripe faint, cream; lateral stripe 
wide, cream; marked with various spots and 
patches of cream. Cream dorsally with a 
few faint mid-dorsal patches of brown. Pro- 
thorax with SD and L setal groups on 
brown pinacula. Thoracic segments 2 and 3 
with seta SDl and setae LI and L3 on 
brown pinacula. Legs cream. Underside 
cream. 

Abdomen (Fig. 3): Mid-dorsal quadrate 
marks light brown, less distinct on seg- 
ments 4-6 and 9; dorsolateral band cream; 
lateral band broad, brown, broken by series 
of cream spots along dorsal margin; distinct 
cream spots slightly dorsal and posterior to 
spiracles. Dl seta inside, D2 seta outside 
dorsal patch. Segments 1 and 2 with SDl, 
LI, and L3 on brown pinacula. Segments 
3-6 and 8 with SDl and L group setae on 
brown pinacula. Prolegs cream; crochets in 
an uniordinal mesoseries. Segments 1 and 
2 with 2 SV setae, SVl represented by a 
sclerotized ring near SV3. 

Pupa 

Diagnosis. — Labial palpus present, con- 
tinuous; profemur present. 

Description. — Male (Figs. 34-36).- 
Length 15.8 ± 1.04 mm (range, 15.6-19.6 
mm) (n = 4). Labial palpi present, elongate, 
continuous. Profemur present. Mesothorac- 
ic leg reaching eye. Wings do not extend 
beyond caudal margin of segment 4. Seg- 
ments 1-3 and 8 with shallow circular pits 
on dorsum; segments 4-7 with shallow cir- 
cular pits in a dense band extending about 
Va width of segment caudally and complete- 



308 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




27 



28 



29 



Figs ^5-^9 Eidepidotis merricki pupa. 25. 3 ventral view. 26. 6 dorsal view (scale = 1.0 mm). 27, S 
ventral view of terminal segments. 28. 9 ventral view of terminal segments. 29. Dorsal view ot ^enmnal segmen s 
(scale = 0.5 mm). Abbreviations: ant =antenna; ao = anal opening; go = genital openmg; Ip - labial palpi. 11 
= prothoracic leg; 12 = mesothoracic leg; mx == maxilla. 



VOLUME 101, NUMBER 2 



309 






32 




Figs. 30-33. Eiilepidotis jiuicida pupa. 30. S ventral view. 31. d dorsal view (scale = 1.0 mm). 32. S 
ventral view of terminal segments. 33, 9 ventral view of terminal segments (scale = 0.5 mm). Abbreviations: 
fl = femur of prothoracic leg; Ip = labial palpi. 



310 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 







Figs M-M Eulepidous supenor pupa. 34. 6 ventral view. 35. S dorsal view (scale - 1.0 mm). 36. 6 
ventral Jiew ol' terminal segments. 37, 9 ventral view of terminal segments (scale = 0.5 mm). Abbreviations: 
fl = femur of prothoracic leg; Ip = labial palpi. 



VOLUME 101, NUMBER 2 



311 



ly encircling segments 5-7; segment 9 
smooth. Genital opening on an oval plate. 
Anal opening caudad of genital opening. 
Cremaster consisting of a large median pair 
of curved hooks and 3 pairs of small hooks, 
1 pair located mediodorsally, 2 pairs locat- 
ed laterally (Fig. 36). 

Female (Fig. 37): Similar to male except: 
length 16.1 ± 1.03 mm (range, 15.0-17.0 
mm) (n=3). Genital opening at caudal border 
of segment 8, dividing segment 8 ventrally. 
Anal opening well caudad of genital opening. 

Host. — Quararibea asterolepis Pitt.; 
family Bombacaceae. 

Natural History. — During late May 
through early June 1985, a massive, highly 
synchronized outbreak of larvae was ob- 
served on the central plateau of Barro Col- 
orado Island, Republic of Panama (Wong et 
al. 1990). The larval host plant was under- 
going leaf flush and the larvae were feeding 
on young leaves and often defoliating the 
trees. Once feeding was completed, the lar- 
vae descended on silk lines and pupated be- 
neath dried leaves on the forest floor. Lar- 
vae were so abundant that it was difficult 
to avoid walking into their silk lines or 
stepping on larvae and pupae. 

A total of 10 pupae and 3 larvae were 
brought to one of us (A A) by Maria Wong, 
Seiji Tanaka, and Peter Becker. Three pupae 
died, and seven were reared to adults. Adult 
no. 1(9) eclosed 21 June from a pupa col- 
lected 7 June. Aduh nos. 4 (d) and 5(9) 
eclosed 23 and 24 June, respectively, from 
pupae collected 1 1 June. Adult nos. 6-9 
eclosed 18 June (9), 20 June (d), and 21 
June {26 S), respectively, from pupae col- 
lected 14 June. All eclosions took place in 
early evening, some as early as 5:30 PM, 
and others as late as 9 PM. 

Discussion. — Adults of E. superior are 
the largest (forewing length 13-9.5 mm) of 
the brown species of Eulepidotis and cannot 
be confused with any other species in the 
genus. It is distributed from Mexico to Pan- 
ama in Central America, northwestern South 
Americain Venezuela, Colombia, and Ecua- 



dor, and in the Caribbean on the islands of 
Puerto Rico, Grenada, and St. Lucia. 

Acknowledgments 
We thank Robin B. Foster of the Smith- 
sonian Tropical Research Institute, Balboa, 
Republic of Panama, for identifying the 
host plant of E. jiincida; Robert W Poole 
of Entomological Information Services, 
Rockville, Maryland, for identifying E. su- 
perior; and Lionel Pagan of U.S.D.A., 
A.PH.I.S., RRQ., San Juan, Puerto Rico, 
for supplying the immature stages and 
reared adults of E. merricki. We thank Wil- 
liam E. Miller, University of Minnesota, St. 
Paul, Minnesota, and Douglass R. Miller 
and David R. Smith of the Systematic En- 
tomology Laboratory, U.S.D.A., Beltsville, 
Maryland, and Washington, D.C., for criti- 
cally reviewing and offering suggestions 
that greatly improved the manuscript. Linda 
Lawrence, Systematic Entomology Labo- 
ratory, U.S.D.A, Washington, D.C., pre- 
pared the habitus illustrations. 

Literature Cited 

Adams, C. D. 1972. Flowering plants of Jamaica. Uni- 
versity of the West Indies, Mona, Jamaica, 848 pp. 

Franclemont, J. G. and E. L. Todd. 1983. Noctuidae, 
pp. 120-159. //; Hodges, R. W. et al., eds.. Check 
list of the Lepidoptera of America North of Mex- 
ico. The Wedge Entomological Research Eoun- 
dation, Washington, D.C., 284 pp. 

Holland, W. J. 1902. Description of an apparently new 
species of Palindia. Entomological News 13: 
172-173. 

Nascimento, M. T. and J. Proctor. 1994. Insect defoli- 
ation of a monodominant Amazonian rainforest. 
Journal of Tropical Ecology 10: 633-636. 

Poole, R. W. 1989. Lepidopterorum Catalogus (New 
Series), Fascicle 118, Noctuidae. E. J. Brilland 
Flora and Fauna Publications, Leiden, 1,313 pp. 

Poole. R. W. and P Gentili (eds.). 1996. Nomina In- 
secta Nearctica. A check list of the insects of 
North America, Vol. 3: Diptera, Lepidoptera, Si- 
phonaptera. Entomological Information Services, 
Rockville, Maryland, 1,143 pp. 

Proctor, G. R. 1984. Flora of the Cayman Islands. Roy- 
al Botanical Gardens, Kew, London, 834 pp. 

Wong, M., J. Wright, S. P Hubbell. and R. B. Foster. 
1990. The spatial pattern and reproductive con- 
sequences of outbreak defoliation in Quararibea 
asterolepis. a tropical tree. Journal of Ecology 78: 
579-588. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999, pp. 312-315 

OBSERVATIONS ON THE PREY AND NEST CLUSTERS OF PODALONIA 
VALIDA (CRESSON) (HYMENOPTERA: SPHECIDAE) 

Kelvin M. O'Neill and Howard E. Evans 

(KMO) Department of Entomology, Montana State University, Bozeman, MT 59717, 
U.S.A. (e-mail: koneill@montana.edu); (HEE) Department of Entomology, Colorado State 
University, Fort Collins, CO 80523, U.S.A. 



Abstract. — In northern Colorado, Podalonia valida preyed upon mature larvae of the 
saltmarsh caterpillar, Estigmene acrea (Drury), the same prey it uses in Arizona. Prey 
were deposited in shallow, single-celled nests that occurred within clusters probably pro- 
visioned by single females. Because females forage for abundant nearby prey, stock each 
nest cell with a single caterpillar, and dig simple nests in a localized area, they have the 
potential to provision multiple nests over short time periods. Several observations are 
consistent with this hypothesis. First, within clusters of nest cells, there was a high degree 
of synchrony in developmental stages of the wasps, suggesting that eggs were laid in 
quick succession. Second, dissections showed that female P. valida carry more mature 
eggs in their ovaries than is typical for non-parasitoid aculeate wasps. 

Key Words: Digger wasp, Sphecidae, nesting behavior, egg size, Estigmene acrea, salt- 
marsh cateipillar, Arctiidae 



Wasps of the genus Podalonia dig short 
burrows terminating in a cell provisioned 
with a single prey (Bohart and Menke 
1976). Podalonia luctuosa (Smith) 
(O'Brien and Kurczewski 1982), Podalonia 
robiista (Cresson) (Kurczewski et al. 1992), 
and Podalonia argentifrons (Cresson) 
(O'Brien 1983) restrict themselves to cut- 
worms (Noctuidae), but may take a variety 
of species at any one site. Similarly, Po- 
dalonia occidentalis Murray apparently 
prey solely on tent caterpillars (Lasiocam- 
pidae: Malacosoma) (Evans 1987) and Po- 
dalonia valida (Cresson) limit themselves 
to arctiid moth cateipillars. Steiner (1974, 
1975) found P. valida specializing on salt- 
marsh caterpillars {Estigmene acrea (Dru- 
ry)) in southern Arizona, whereas Rust et 
al. (1985) found two female P. valida tak- 
ing Apantesis proxima (Guerin-Meneville) 
on San Clemente Island, California. Here, 



we present results of our observations of 
the nesting behavior and prey of P. valida 
in northern Colorado and compare our find- 
ings to those of Steiner, who concentrated 
on the hunting and territorial behavior of 
females. 

Methods 

We studied P. valida females at the Paw- 
nee National Grasslands in northern Weld 
Co., Colorado, from 12 July to 13 August 
1984. The site was situated along a sandy, 
little-used road that passed through prairie 
consisting of grasses mixed with several 
common forbs, notably white sweetclover 
(Melilotus alba Desr.) and sunflower (He- 
li an thus spp.). 

Results and Discussion 

Our 64 prey records indicate that P. val- 
ida preyed exclusively on mature larvae of 



VOLUME 101, NUMBER 2 



313 



the saltmarsh caterpillar Estigmene acrea 
(Drury) (Arctiidae), the same species used 
by this wasp in Arizona (Steiner 1974). The 
prey of P. valida, which were probably tak- 
en on their host plants (white sweetclover), 
were carried in the mandibles as females 
walked forward straddling the prey. One fe- 
male carried a caterpillar at least 10 m be- 
fore reaching her nest. Prey-carrying fe- 
males occasionally stopped and cached 
their prey —5 cm above the soil surface on 
small plants, where they were left for sev- 
eral minutes while females searched for 
their nests. 

Unlike other species of Podalonia, P. 
valida dig nests before hunting (Steiner 
1974, 1975). Each nest at our site consisted 
of a short, oblique burrow about 1 cm in 
diameter, terminating in a single 1 X 3 cm 
cell situated 3-5 cm beneath the soil surface 
(N = 7). Typically, when digging the nest, 
a female backed out 5-10 cm from the bur- 
row entrance and scraped soil backwards 
with her forelegs, while elevating her ab- 
domen, flicking her wings rapidly, and 
buzzing loudly. Upon returning to her nest 
with prey, a female dropped it at the en- 
trance, cleared the burrow, and pulled the 
prey inside while moving backwards. After 
several minutes inside the nest, during 
which time she laid an egg on the prey, the 
female emerged and permanently closed the 
bunow by scraping in soil from the edge of 
the burrow and placing lumps of soil or 
pebbles in the hole. She then tamped the 
loose soil with her head (mandibles wide 
open) or with a lump of soil which was held 
in the mandibles and which broke up due 
to the impact. The entire sequence between 
entering the hole with prey and completing 
closure typically took about 4 minutes. 

These observations are in accord with 
those of Steiner (1974, 1975), who found 
that individual females provisioned series 
of single-celled nests within small patches 
of bare soil sometimes no more than 60 cm 
across. Although we did not observe indi- 
vidually marked females over prolonged 
periods, we also found that cells tended to 



be clustered. In 2 of our 7 excavations of 
recently completed nests, we unearthed just 
a single cell, but in the others, we found 
clusters of 8, 15, 16, 18, and 27 cells within 
areas no more than 0.25 m-. Some cells 
were separated by as little as 5 cm, but were 
definitely parts of different nests. Clusters 
of nest cells were well-separated from one 
another and each was apparently used ex- 
clusively by a single female, who vigorous- 
ly attacking conspecific females intruding 
upon their nest cluster in interactions that 
included bouts of grappling between the 
combatants. Similar interactions between P. 
valida females have been described in de- 
tail by Steiner (1975). 

The contents of unparasitized and non- 
moldy P. valida cells within clusters sug- 
gest that, if a single female was responsible 
for all of the cells, some were provisioned 
during short time intervals. For example, in 
the cluster of 15 cells, 5 had prey with wasp 
eggs, 3 had prey with small wasp larvae, 
and 1 had a large wasp larva. In the cluster 
of 7 cells, all had prey with unhatched eggs. 
Similarly, the cluster of 18 cells included 7 
prey with wasp eggs and 7 with small wasp 
larvae, and the cluster of 16 cells contained 
7 with eggs or small larvae. 

The potential ability of P. valida fe- 
males to provision multiple nests in rapid 
succession may be possible because they 
1 ) forage for abundant prey nearby. 2) 
stock each cell with a single prey, and 3) 
dig simple, shallow nests without search- 
ing widely for successive nesting sites. 
This strategy may allow them to exploit a 
single developmental stage of a single prey 
species that is available for just a brief pe- 
riod during the summer However, the rap- 
id stocking of multiple nests would also 
require that a female produce the requisite 
numbers of eggs. Typically, female sphe- 
cids carry no more than two mature eggs 
at a time, although they have three ovari- 
oles in each ovary (Iwata 1964, O'Neill 
1985). Nevertheless, the three P. valida fe- 
males that we captured within minutes of 
their laying eggs had 3, 5, and 6 mature 



314 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



(or nearly mature) oocytes still in their 
ovaries. Thus, the latter female had been 
carrying seven well-developed oocytes just 
prior to capture and had one ovariole with 
two mature oocytes. In Iwata's extensive 
survey of the ovaries of solitary aculeate 
wasps, only parasitoid sphecids of the gen- 
era Larra (with up to 21 mature oocytes) 
and Chlorion (with up to 10) carried more 
mature eggs than the maximum for P. val- 
ida. Such high levels of short-term fecun- 
dity (for a digger wasp), however, may be 
achieved at the cost of producing smaller 
eggs. The mature oocytes of P. valida 
ranged in length from 2.6-3.1 mm in 
length and one egg found on prey was 2.9 
mm long. In contrast, in the similarly-sized 
but less fecund wasp Philcmthus hicinctus 
(Mickel), mature oocytes and newly laid 
eggs ranged as high as 5.4 mm in length, 
and had perhaps 3-4 times the volume of 
P. valida eggs (O'Neill 1985). Bemhecinus 
quinqiiespinosus (Say) and Philanthus 
pulcher Dalla Torre carry 1-2 mature oo- 
cytes within the same size range as P. val- 
ida, although they are both much smaller 
wasps. 

Of the 86 cells we excavated, four had 
fly puparia 0.5-1.0 cm below the cell, 
which in each case-contained the remains 
of a single saltmarsh caterpillar. One pu- 
parium gave rise to a tachinid of the genus 
Exorista, at least one species of which is a 
known parasitoid of E. acrea (Arnaud 
1978). The wasps had probably captured 
previously parasitized caterpillars and suf- 
fered incidental cleptoparasitism. Another 
23% of the cells contained molded or rot- 
ting caterpillars. 

Our observations complement those of 
Steiner (1974, 1975), indicating that two 
widely separated populations of P. valida 
have identical prey preferences and simi- 
lar nesting behaviors that are unique for 
this genus. Podalonia valida seems to 
have adopted a strategy intermediate be- 
tween those of the relatively high fecun- 
dity parasitoid sphecids {Larra and Chlo- 
rion) and the more typical low fecundity 



nest provisioning species that spend con- 
siderable time and energy on each off- 
spring. This strategy is facilitated by the 
greater number of mature eggs they carry 
and, perhaps, by shorter periods of search- 
ing for potential nest sites once nesting 
has begun. 

Acknowledgments 

We thank Patricia Denke, Wendell Mor- 
rill, Mark O'Brien, and Catherine Seibert 
for commenting on the manuscript and N. 
Woodley (Systematic Entomology Labora- 
tory, USDA, Washington, DC) for deter- 
mination of the tachinid fly. The work was 
supported by the National Science Foun- 
dation, the Colorado Agricultural Experi- 
ment Station, and and the Montana Agri- 
cultural Experiment Station. 

Literature Cited 

Arnaud, P. H. 197S. A host-parasite catalog of North 
American Tachinidae (Diptera). Miscellaneous 
Publication No. 1319, Science and Education Ad- 
ministration, United States Department of Agri- 
culture, 860 pp. 

Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps 
of the World: A Cieneric Revision. Berkeley: Uni- 
versity of California Press, ix + 96.'i pp. 

Evans, H. E. 1987. Observations on the prey and nests 
of Podalonia occidentalis Murray (Hymcnoptera: 
Sphecidae). Pan-Pacific Entomologist 63: 130- 
134. 

Iwala, K. 1964. Egg gigantism in subsocial Hymcnop- 
tera, with ethological discussion on tropical bam- 
boo carpenter bees. Nature and Life in .Southeast 
Asia 3: 399-434. 

Kurczewski, F. E., M. E O'Brien, and M. G. Spofford. 
1992. Nesting behavior of Podalonia rohusta 
(Cresson) (Hymenoptera: Sphecidae). Journal of 
Hymenoptera Research 1 : 235-239. 

O'Brien, M. F 1983. Observations on the nesting be- 
havior of Podalonia argentifrons. Southwestern 
Entomologist 8: 194-197. 

O'Brien, M. E and E E. Kurczewski. 1982. Ethology 
and overwintering of Podalonia luctiiosa (Hy- 
menoptera: Sphecidae). Great Lakes Entomologist 
15: 261-275. 

O'Neill, K. M. 1985. Egg size, prey size, and sexual 
size dimoiphism in digger wasps (Hymenoptera: 
Sphecidae). Canadian Journal oi Zoology 63: 
2187-2193. 

Rust, R., A. S. Menke, and D. Miller. 1985. A biogeo- 
graphic comparison of the bees, sphecid wasps. 



VOLUME 101, NUMBER 2 



315 



and mealybugs of the California Channel Islands 
(Hymenoptera, Homoptera). pp. 29-59 in Menke, 
A. S. and D. R. Miller, eds.. Entomology of the 
Channel Islands. Santa Barbara Museum of Nat- 
ural History, Santa Barbara, California. 
Steiner, A. L. 1974. Unusual caterpillar-prey records 
and hunting behavior for a Podalonio digger 



wasp: Podalonia valida (Cresson). Pan-Pacific 
Entomologist 50: 13-11. 
Steiner, A. L. 1975. Description of territorial behavior 
of Podalonia valida (Hymenoptera: Sphecidae) 
females in southeast Arizona, with remarks on 
digger wasp territorial behavior. Quaestiones En- 
tomologica 11: 113-137. 



PROC. ENTOMOL. SOC. WASH. 

101(2), 1999, pp. 316-324 

IDENTIFICATION OF LATE-INSTAR NYMPHS OF 
COCKROACHES (BLATTODEA) 

Kevin T. Hathorne and Patricia A. Zungoli 

Department of Entomology, Clemson University, Box 0365, Clemson, SC 29634-0365, 
U.S.A. (e-mail: kevinh@clemson.edu) 



Abstract. — A dichotomous key to late-instar nymphs of 12 cockroach species common 
in the United States is presented. Notes on biology, distribution, and taxonomy are given. 

Key Words: Insecta, Blattodea, late-instar nymphs, descriptions, dichotomous key 



More than 3600 species of cockroaches 
are known worldwide, 69 of which are 
found in North America. Fifteen to twenty 
of these species are of economic importance 
as nuisance pests. Classification of cock- 
roaches has undergone many changes. Rehn 
(1951) classified adult cockroaches based 
on wing characteristics. Princis (1960) pro- 
vided a comprehensive history of cock- 
roach taxonomy and reported previously 
unknown information on cockroach evolu- 
tion. McKittrick (1964) grouped the cock- 
roaches with mantids based on evolutionary 
studies, morphology of genitalia and pro- 
ventriculus, and oviposition behavior. Corn- 
well (1968) provided a thorough review of 
the history of cockroach classification. He 
took the reader from 1758 when all cock- 
roaches were placed in the genus Blatta, in 
the order Coleoptera, through the revision 
of Imms' Textbook of Entomology (Rich- 
ards and Davies 1957) in which cockroach- 
es are placed in their own order, Dictyop- 
tera. Cockroaches are currently placed in 
the order Blattodea as recognized by the 
Entomological Society of America (Bosik 
1997). 

Dichotomous keys to the cockroaches are 
primarily restricted to the adult stage 
(Blatchley 1920, Rehn 1950, Pratt and Sto- 
janovich 1962, Dakin and Hays 1970) ex- 



cept for those by Powell and Robinson 
(1980), Fisk (1987), and Gordon (1992). 
Powell and Robinson (1980) included only 
first-instar nymphs of five Periplaneta spe- 
cies and Gordon (1992) distinguished the 
mid-instar nymphs of Periplaneta ameri- 
cana and P. fuliginosa. Fisk (1987) includ- 
ed nymphs of 16 cockroach species with 
identification based on comparative char- 
acters. 

Because behavior and habitat preference 
differ substantially among species, and de- 
velopment of effective control strategies de- 
pends on correct identification, a dichoto- 
mous key to late-instar cockroach nymphs 
is needed. The present work provides a 
means of identifying 1 2 of the pest species 
found in North America. 

Materials and Methods 

Based on their pest status, late-instar 
nymphs of 12 species of cockroaches rep- 
resenting 3 families (Blaberidae, Blattelli- 
dae, and Blattidae) are included in this key. 
A cockroach may go through 5 to 12 molts 
before reaching the adult stage. The indi- 
vidual specimens were selected for size (ap- 
proximately the same size as the adult of 
the same species). Exact instar was not 
known for each individual. Specimens of 
the following eight species were obtained 



VOLUME 101. NUMBER 2 



317 



from colonies maintained by the Clemson 
University Urban Entomology Laboratory: 
Blattella germanica (L.), Supella longipal- 
pa (E), Pehplaneta americana (L.), Peri- 
planeta australasiae (E), Pehplaneta briin- 
nea Burmeister, Periplaneta fuliginosa 
(Serville), Blatta ohentalis (L.), and Par- 
coblatta lata (Brunner). Three species 
{Blattella asahinai Mizukubo, Panchlora 
nivea [L.], and Eurycotis floridana [Walk- 
er]) were obtained from the USDA-ARS 
laboratory in Gainesville, Florida. One spe- 
cies, Blattella vaga Hebard, was obtained 
from Virginia Polytechnic Institute and 
State University, Blacksburg, Virginia. 

Ten to twenty specimens of each species 
were examined for morphological differ- 
ences, using a dissecting microscope 
(WILD Heerbrugg Switzerland M5-3984). 

Three species of cockroaches in this key 
are wingless as adults. These wingless spe- 
cies, on close examination, have truncated 
wings (B. orientalis (female), E. floridana), 
or wing pads (female P. lata). This char- 
acter may confuse the identification of the 
adult form with a late-instar nymph of an- 
other species. All characters used in this 
key can be seen when the specimen is 
viewed dorsally. 

Key to Late-Instar Cockroach Nymphs 

1. Body tan. Thorax with 2 dark brown to black 
parallel lines 2 

- Body color variable. Thorax without parallel 
lines 4 

2. Abdomen with 2 tan dots on some or all ter- 
gites in center of dark longitudinal area. Ab- 
domen with tan markings on lateral edges 
separated by dark brown markings between 
tergites (note: All above characters may vary 
slightly or may not be present) 

Blattella gemuinica (Fig. 2) 

- Abdomen with markings different from above 3 

3. Abdomen with tan, horizontal bar on some or 
all tergites in center of dark longitudinal area. 
Body with black, vertical line through center. 
Cercus dark brown to black on terminal ends. 
Abdomen with tan markings on lateral edges 
separated by dark brown markings between 
tergites (note: All above characters may vary 
slightly or may not be present) 

Blattella vai>a (Fig. 3) 



- Abdomen with tan, horizontal bar bearing an- 
terior emargination on some or all tergites in 
center of dark longitudinal area. Body without 
black, vetical line. Cercus tan with dark 
brown to black markings on terminal ends. 
Abdomen with tan markings on lateral edges 
not separated by dark brown markings be- 
tween tergites (note: All above characters 
may vary slightly or may not be present) . . 

Blattella asahinai (Fig. 4) 

4. Cercus longer than distance between their ba- 
ses 5 

- Cercus shorter than distance between their ba- 
ses 10 

5. Body variably black and tan. Pronotum with 
black horizontal bar bearing anterior and pos- 
terior median emarginations 

Periplaneta australasiae (Fig. 5) 

- Body uniform reddish-brown or with black on 
lateral edges of abdominal tergites or tan with 
dark brown markings. Pronotum with mark- 
ings different from above 6 

6. Pronotum dark brown with tan on lateral edg- 
es. 2nd and 3rd thoracic sclerites with hori- 
zontal dark brown markings. Length less than 

15 mm Supella longipalpa (Fig. 1) 

- Pronotum with markings different than above. 
2nd and 3rd thoracic sclerites without hori- 
zontal dark brown markings. Length greater 
than 15 mm 7 

7. Pronotum dark with little color variation. 
Male with styli shorter than 10th tergite . . . 

Periplaneta fuliginosa (Fig. 6) 

- Pronotum with some color variation, dark- 
ened areas. Styli variable 8 

8. Abdomen black, with lighter areas in center 
of tergites. 10th tergite truncate 

Parcoblatta lata (Fig. 7) 

- Abdomen with color pattern different from 
above. 10th tergite notched 9 

9. Male with styli slightly longer than distance 
between their bases. Cercus with last segment 
twice as long as wide (Fig. 8a) 

Periplaneta americana (Fig. 8) 

- Male with styli shorter than distance between 
their bases. Cercus with last segment less than 
twice as long as wide (Fig. 9a) 

Periplaneta hritnnea (Fig. 9) 

10. Body black. Cercus longer than 10th tergite 

Blatta orientalis (Fig. 10) 

- Body light brown or dark red and black. Cercus 

as long as, or shorter than, 10th tergite .... 11 
1 1. Body dark red in center, with black laterally. 
9th abdominal tergite with posterior corners 
prolonged backward into sharp points. Tho- 
racic and abdominal lateral margins forming 
a smooth line in dorsal view. 10th tergite 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



shorter than cercus 

Eitrycotis fioridaiia (Fig. 1 1 ) 

- Body uniformly reddish brown. 9th abdomi- 
nal tergite without points. Thoracic and ab- 
dominal lateral margins not forming a smooth 
line in dorsal view. 10th tergite longer than 
cercus Panchlora nivea (Fig. 12) 

Notes on Species 

Supella longipalpa (Fig. 1). — The 
brown-banded cockroach nymph has a 
broad black stripe on the pronotum and oth- 
er black markings on the thorax. It is typi- 
cally less than 1 5 mm. This species is found 
throughout the United States in relatively 
sanitary conditions such as office buildings. 
It prefers temperatures over 27°C and is of- 
ten found above floor level. It will infest 
furniture, and deposit egg cases behind pic- 
ture frames and appliances. 

Blattella germanica, B. asahinai, and B. 
vaga are similar species. The German, 
Asian, and field cockroach nymphs are tan 
with two broad, dark brown to black lon- 
gitudinal stripes on the thorax. Because 
these species are so similar to one another, 
multiple characters must all be used for 
identification of species. As late-instar 
nymphs there are differences in body size 
between these three species. However, these 
size variations may overlap. 

Blattella germanica (Fig. 2). — When 
body length is compared to the other two 
Blattella species represented in this key, it 
usually is the median. The German cock- 
roach is perhaps the most economically im- 
portant of all cockroach pests. It is found 
throughout the United States in human hab- 
itations, and rarely outside. 

Blattella vaga (Fig. 3). — The field cock- 
roach nymph is typically smaller and lighter 
in color than B. germanica and B. asahinai. 
It is found in the southwestern United 
States in irrigated fields and yards. 

Blattella asahinai (Fig. 4). — The Asian 
cockroach nymph is typically 2 to 3 mm 
longer than B. germanica. Most individuals 
have three small black dots arranged in a 
triangle, a character not always observed in 
B. germanica or B. vaga. Found primarily 



outdoors in Florida, it occasionally enters 
structures. 

Parcoblatta lata (Fig. 7). — -The broad 
wood cockroach nymph has a uniformly 
reddish brown thorax. The abdomen is 
mostly black with lighter markings in the 
center of the first three to four tergites. The 
10th tergite is pointed and unnotched. It is 
found throughout the United States in 
wooded habitats. Adult females are wing- 
less. Adult males will occasionally enter 
buildings. 

Periplaneta australasiae (Fig. 5). — The 
Australian cockroach nymph is easily dis- 
tinguished by its coloration, it is black and 
tan and has a dark horizontal bar with an- 
terior and posterior emarginations on the 
pronotum. The adults share this character. 
This species is most commonly found in 
both indoor and outdoor situations in the 
coastal states of the southern United States 
and portions of California. 

Periplaneta fuliginosa (Fig. 6). — -The 
smokybrown cockroach nymph, very simi- 
lar to P. americana and P. brunnea, can be 
distinguished by its uniform reddish brown 
color. It is found in the southeastern United 
States in hot, humid areas in and around 
structures. 

Periplaneta americana (Fig. 8). — Like 
P. brunnea, the American cockroach 
nymph is reddish brown with variable dark- 
ened areas. The cerci are long and slender 
and the last segment is twice as long as it 
is wide. If styli are present, they are longer 
than the distance between their bases. This 
species is found throughout the United 
States in association with buildings and 
warm, humid areas. 

Periplaneta brunnea (Fig. 9). — The cerci 
and styli on the brown cockroach are the 
key characters used to distinguish this spe- 
cies from P. americana. The cerci are more 
flattened and broader and the last segment 
is not twice as long as it is wide. If styli 
are present, they are shorter than the dis- 
tance between their bases. The brown cock- 
roach is found around buildings in the 
southeastern United States. 



VOLUME 101. NUMBER 2 



319 




Figs. 1-2. Dorsal view of late-instar nymphs. l.Supella longipalpa, the brown-banded cockroach. 2. Blatella 
?rmaiiica, the Gerinan cockroach. 



320 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





Figs. 3-4. Dorsal view of late-instar nymphs. 3. Blatella va.uci. Held coctcroach. 4. B. asahinai. the Asian 
cockroach. 



VOLUME 101. NUMBER 2 



321 




Figs. 5-7. Dorsal view of late-instar nymphs. 5, Periplanetci austndusiae. the Australian cockroach, 6. P. 
ftiliglwsci. the smokybrown cockroach. 7, Parcohlattu lata, the broad wood cockroach. 



322 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 8-9. Dorsal view of late-instar nymphs. 8. Periplaneta ameiiccma, the American cockroach; 8a - 
dorsal view of left cercus. 9. P. hrunuea. the brown cockroach; 9a = dorsal view of left cercus, note size and 
shape of last segment. 



VOLUME 101. NUMBER 2 



323 




Figs. 10-12. Dorsal view of late-instar nymphs. 10, Blatta orientalis, the Oriental cockroach. 1 1. Eiuycotis 
ftoridana, the Florida woods cockroach. 12, Panchlora nivea, the Cuban cockroach. 



Blatta orientalis (Fig. 10). — The Oriental 
cockroach, easily identified by its uniform 
dark, reddish-black color, can also have a 
shiny appearance. Adult females have trun- 
cated wings. It is found in and around struc- 
tures over most of the temperate United 
States and Canada. 

Eurycotis floridana (Fig. 1 1). — The Flor- 
ida woods cockroach nymph is compara- 
tively large (35 mm) and dark in color. The 
center of the body is dark red and the outer 



edges are black. The posterior comer of the 
9th abdominal tergite is prolonged back- 
ward to sharp points. Adults have truncated 
wings. An important pest in Florida, it also 
is found in southeastern Georgia and along 
the lower Gulf and Atlantic coasts of the 
United States. The adult emits an oily liquid 
with a repellent odor. 

Panchlora nivea (Fig. 12). — The Cuban 
cockroach nymph is reddish brown, and the 
lateral margins of the thoracic and abdom- 



324 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



inal segments appear jagged when viewed 
dorsally. The 10th tergite is rectangular and 
longer than the cerci. It is found around 
houses and wooded areas throughout Flor- 
ida and along the Gulf Coast of the United 
States. 

Acknowledgments 

We thank Peter H. Adler (Clemson Uni- 
versity, Clemson, South Carolina) for his 
advice with this project. We also thank 
Mary Ross (Virginia Polytechnic Institute 
and State University, Blacksburg, Virginia), 
Richard Brenner (USDA-ARS MAVERL 
Laboratory, Gainesville, Florida), and Phil 
Koehler (University of Florida, Gainesville, 
Florida) for generously providing us with 
cockroaches. Al Wheeler and Mike Hood, 
both of Clemson University (Clemson, 
South Carolina), reviewed and improved 
this manuscript. This manuscript is techni- 
cal contribution number 4399 of the South 
Carolina Agricultural Experiment Station. 

Literature Cited 

Blatchley, W. S. 1920. Orthoptera of Northeastern 
America. Nature Publishing Company, Indianap- 
olis. 784 pp. 

Bosik, J. B. 1997. Common Names of Insects & Re- 
lated Organisms 1997. The Entomological Society 
of America. Lanham, Maryland. 232 pp. 

Cornwell, R B. 1968. The Cockroach Vol. 1. A Lab- 



oratory Insect and an Industrial Pest. Hutchinson 
& Co. London. 391 pp. 

Dakin. M. E.. Jr and K. L. Hays. 1970. A Synopsis of 
Orthoptera (sensu lato) of Alabama. Auburn Uni- 
versity Agricultural Experiment Station Bulletin 
404, 118 pp. 

Fisk, F W. 1987. Order Blattodea, pp. 120-131. In 
Stehr, F W., ed.. Immature Insects. Kendall/Hunt 
Publishing, Dubuque, Iowa. 

Gordon, J. M. 1992. Comparison of mid-instar nymphs 
of Periplcmeta fuliginosa (Serville) and Peripla- 
netci americana (L.) (Blattodea: Blattidae) Pro- 
ceedings of the Entomological Society of Wash- 
ington 94: 151-156. 

McKittrick, F A. 1964. Evolutionary Studies of Cock- 
roaches. Cornell University Agricultural Experi- 
ment Station. Memoir # 389, 197 pp. 

Powell, P. K. and W. H. Robinson. 1980. Descriptions 
and keys to the first-instar nymphs of five Peri- 
pluneta species (Dictyoptera: Blattidae). Proceed- 
ings of the Entomological Society of Washington 
82: 212-228. 

Pratt, H. D. and C. J. Stojanovich. 1962. Key to the 
Common Cockroaches found in the United States 
of Public Health Importance. United States De- 
partment of Health and Environmental Welfare. 
Public Health Service. Centers for Disease Con- 
trol. 

Princis, K. 1960. Zur Systematik der Blattarien. Eos 
36: 429-449. (Authors have not read original 
work.) 

Rehn. J. W. H. 1950. A key to the genera of North 
American Blattaria, including established adven- 
tives. Entomological News 61(3): 64-67. 

. 1951. Classification of the Blatteria as indi- 
cated by their wings (Orthoptera). Memoirs of the 
American Entomological Society 14: 1-134. 

Richards. O. W. and R. G. Davies. 1957. A. D. Imms. 
A General Textbook of Entomology. Ninth Edi- 
tion. Methuen & Co. Ltd. London. 886 pp. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999. pp. 325-331 

STUDIES ON STONEFLIES OF NORTH DAKOTA WITH THE 
DESCRIPTION OF A NEW PERLESTA SPECIES (PLECOPTERA: FERLIDAE) 

Boris C. Kondratieff and Richard W. Baumann 

(BCK) Department of Bioagricultural Sciences and Pest Management, Colorado State 
University, Fort Collins, CO 80523, U.S.A. (e-mail: bkondrat@ceres.agsci.colostate.edu); 
(RWB) Department of Zoology and Monte L. Bean Life Science Museum, Brigham 
Young University, Provo, UT 84602, U.S.A. (e-mail: richard_baumann@byu.edu) 



Abstract. — Records for fifteen stonefly taxa, including eleven confirmed species are 
given for North Dakota. Literature records for an additional three species could not be 
confirmed. Perlesta dakota, n. sp., is described from the adult male, female and egg. 
Diagnostic characters are presented using illustrations and a SEM photomicrograph. 

Key Words: stoneflies, Plecoptera, North Dakota, Perlesta, new species 



Only two species of stoneflies were listed 
for North Dakota by Stark et al. (1986), a 
state covering 18,299,503 hectares that in- 
cludes two major river systems, the Mis- 
souri and the Red River of the North. The 
landscapes of North Dakota, as common 
with most Northern Great Plains states, ex- 
hibit relatively little topography, and the 
state is better known for its numerous prai- 
rie marshes (Van der Valk 1989). Many 
counties of the state have 80-90% of their 
total area in agricultural production. 

The Red River forms the eastern border 
of the state with Minnesota, traversing the 
former bed of the glacial Lake Agassiz. The 
area west of the Red River Valley, a region 
often referred to as the Drift Prairie is poor- 
ly drained, with few streams of any gradi- 
ent. West of the Drift Prairie is the Missouri 
Plateau, also an area of poor drainage, and 
further west is the Missouri River and its 
associated tributaries. These streams are of- 
ten located in broad valleys of native grass- 
es or hay and small grain production areas. 
The Little Missouri River is a prominent 
feature of western North Dakota and drains 
through the spectacular landscape known as 



the "Badlands." Streams of the Badlands 
are mostly silty and relatively unproductive 
(Gordon and Post 1965). A southern exten- 
sion of the Northern Coniferous Forest can 
be found in the northwestern portion of the 
state. 

Despite the presence of suitable lotic 
habitats throughout the state, only nine spe- 
cies of stoneflies have been recorded from 
North Dakota. Stark et al. (1986) in their 
compilation of North American stonefly 
records, listed only Strophopteryx fasciata 
(Burmeister) and Haploperla orpha (Pri- 
son). However, we have found seven addi- 
tional records in the literature. Harden and 
Mickel (1952) noted a record for Isoperla 
longiseta (Banks), and Neel (1985) in his 
ecological study of the Turtle River in east- 
ern North Dakota discussed a species of Ac- 
roneuria, Perlesta placida (Hagen), and 
Taeniopteryx nivalis (Fitch). Finally, Stoaks 
(1975) listed nymphal records of Pteronar- 
cys dorsata (Say), P. placida, A. arenosa 
(Pictet) and /. bilineata (Say) from the For- 
est River. 

A recent collecting trip by the authors 
into southwestern North Dakota yielded an 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 1-6. Perlesta dakota. 1, Adult head and pronotum. 2, Male terminalia, dorsal. 3, Paraproct, caudal. 4, 
Paraproct, lateral. 5, Penis. 6, Female subgenital plate, ventral. 



undescribed species of Perlesta. The genus 
Perlesta in North America presently in- 
cludes sixteen species (Stark 1989, Poulton 
and Stewart 1991, Stark and Rhodes 1997, 
Kirchner and Kondratieff 1997, DeWalt et 
al. 1998). The terminology in the descrip- 
tion follows that proposed by Stark (1989). 



Perlesta dakota Kondratieff and 
Baumann, new species 

(Figs. 1-7). 

Male. — Forewing length 8-9 mm. Gen- 
eral color brown. Head yellow with dark 
brown pattern as Fig. 1. Pronotum dark 
brown (Fig. 1 ). Antennal scape and pedicel 



VOLUME 101, NUMBER 2 



327 



brown, basal flagellular segments becoming 
darker brown beyond segments 10-16. Fore- 
femur dorsally brown. Wing membrane and 
veins brown except costal margin pale. Ab- 
dominal terga brown, sterna yellow brown 
usually with darker brown triangle-shaped 
shading. Cercus yellow brown. Tergum 10 
mesal sclerite not divided, sensilla basicon- 
ica small and scattered (Fig. 2). Paraproct 
short in caudal view (Fig. 3), more slender 
in lateral view (Fig. 4), spine anteapical and 
directed mesad (Figs. 2 and 3). Penis tube 
+ sac moderately long with raised shelf 
dorsobasally; caecum prominent, dorsal 
hair patch broad basally narrowing to base 
of caecum, lateral hair patch elongate (Fig. 
5). Female. Forewing length 10-12 mm. 
Color pattern similar to male but paler. Sub- 
genital plate truncate with notch deep and 
narrowly V-shaped (Fig. 6). 

Egg. — Collar button-like. Chorion sur- 
face finely pitted with scattered coarser pit- 
ting. Micropylar row in posterior region 
(Fig. 7). 

Nymph. — Unknown. 

Types. — Holotype d, allotype 9: North 
Dakota, Hettinger Co., Cannonball River, at 
New England, Hwy 22, 15 July 1997, R. 
W. Baumann and B. C. Kondratieff. Para- 
types, same data as holotype: 14 6,9 9; 
Ransom Co., Fort Ransom State Park, 1 1 
July 1970, S. M. Anders, 1 c?, 1 9 
(NDSU); Stark Co., Heart River, Hwy 22, 
Dickinson, 14 July 1997, R. W. Baumann 
and B. C. Kondratieff, 2 9 (BYUC, 
CSUC). Holotype and allotype deposited in 
the National Museum of Natural History, 
Smithsonian Institution, other paratypes at 
the Monte L. Bean Life Sciences Museum, 
Brigham Young University (BYUC); C. R 
Gillette Museum of Arthropod Diversity, 
Colorado State University (CSUC); North 
Dakota State University (NDSU) and the B. 
P Stark Collection (BPSC). 

Etymology. — The Sioux were the largest 
tribe of Native Americans of the North 
American Plains and prairies, and are called 
the Dakota in the Santee dialect. 

Diagnosis. — Perlesta dakota appears 




Fig. 7. Perlesto dakota, scanning electron photo- 
micrograph of egg. 



similar to a group of dark species, including 
P. cinctipes (Banks), P. adena Stark, P.fus- 
ca Poulton and Stewart, and P. xube Stark 
and Rhodes. Perlesta dakota seems most 
similar to P. fusca, a species distributed 
throughout the Ozark-Ouachita Mountain 
region of Arkansas, Missouri and 
Oklahoma (Poulton and Stewart 1991). 
Males of P. dakota can be distinguished 
from P. fusca by the well developed thumb- 
like caecum (Fig. 5) and maculation of the 
head (Fig. 1). The penis structure of P. dak- 
ota is similar to P. decipiens (See Stark 
1989; Figs. 42-43) but the former species 
can be distinguished by the dark brown 
wing and body coloration and head pattern. 
Females of P. dakota can be distinguished 
from P. fusca by the more truncate lobes of 
the subgenital plate and the deep and nar- 
row notch (Fig. 6). The egg collar of P. 
dakota is button-like (Fig. 7), and the cho- 
rion finely pitted (Fig. 8). Poulton and 
Stewart (1991) indicated that the egg of P. 
fusca lacks a collar and the surface of the 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



egg is reticulate. The egg of P. decipiens 
has a distinctive short collar. 

Remarks. — The type locality, the Can- 
nonball River at New England, is typical of 
an agriculturally-impacted stream of the re- 
gion, little or no riparian vegetation, heavily 
silted and with only few small riffles. 
Adults of P. dakota were collected by beat- 
ing tall overhanging streamside grasses. 
Other aquatic insects collected with P. dak- 
ota include the damselfly Calopteryx ae- 
quabilis Say, the mayfly Caenis sp., and the 
caddisflies, Cheiimatopsyche pettiti (Banks), 
Ceratopsyche morosa (Hagen), Hydroptila 
consimilis Morton, and Limnephiliiis hyali- 
nus Hagen. 

North Dakota Stonefly Records 

Collections examined, primarily from 
North Dakota State University (NDSU), 
yielded new records or added to or substan- 
tiated previous records of North Dakota 
stoneflies. Other institutions listed include 
Brigham Young University (BYUC), Col- 
orado State University (CSUC); Dickinson 
State University (DSUC); Minot State Uni- 
versity (MSUM); Mississippi College 
(BPSC); Museum of Comparative Zoology, 
Harvard University (MCZC), University of 
Minnesota (UMSP), and University of Mis- 
sissippi (UMIC). Some of these records are 
only identified to the generic level, but are 
included for distributional purposes, since 
so few stonefly records are currently avail- 
able for North Dakota. A total of fifteen 
stonefly taxa are known from the state of 
North Dakota, representing at least eleven 
species. 

AUocapnia sp. 

Records. — Ransom Co., cold spring-fed 
stream, joining Sheyenne River, Hwy 46, 
Little Yellowstone Park, 1 Nov 1962, R. D. 
Gordon, 6 N (NDSU). 

Several species of this genus of winter 
stoneflies could occur in North Dakota, es- 
pecially A. pygmaea (Burmeister) (Ross 
and Ricker 1971). 



Oemopteryx fosketti (Ricker) 

Records. — Billings Co., Little Missouri 
River, Medora, 19 March 1997, C. R Milne, 
16 d, 6 9 (BYUC, CSUC, DSUC); Little 
Missouri River, Sully Creek Campground, 

19 March 1997, C. P Milne, 11 d, 1 9 
(BYUC). 

The species originally described from 
Saskatchewan, Canada, can be abundant in 
large and often silty rivers of the Missouri, 
Colorado, and Saskatchewan drainages 
(Baumann et al. 1977). Records for this 
species are known from nearby Dawson 
Co., Montana. The males have peculiar up- 
turned forewing tips. 

Strophopteryx fasciata (Burmeister) 

Records. — Cass Co., Fargo, 24 April 
1900, E. Cleveland, 1 S (MCZC). 

Strophopteryx fasciata is a widespread 
species occurring throughout eastern and 
midwestern North America, and typically a 
larger stream species emerging in late win- 
ter and early spring (Stewart and Stark 
1988). 

The record for T. nivalis by Stoaks 
(1975) could not be confirmed due to the 
lack of material, but is possible, since this 
species is common in Minnesota (Ricker 
and Ross 1968). 

Pteronarcys pictetii Hagen 

Records. — North Dakota: Cavalier Co., 

20 Oct 1962, 2 N (NDSU); Pembina Co., 
Pembina River, Walhalla City Park, 1 July 
1970, Perkins and R. L. Post, 1 $ (NDSU); 
Pembina River, Walhalla, 9 July 1961, R. 
L. Post and H. Osborn, 4 N (NDSU); same 
but 20 Oct 1962, E. Saugstad, 2 N (NDSU); 
Richland Co., Sheyenne River, 14 mi NW 
Walcott, 24 June 1963, D. Aarhus, 1 $ 
(NDSU). 

Pteronarcys pictetii has been reported 
from adjacent Minnesota and Manitoba 
(Stark et al. 1986), and was expected in the 
state. The separation of the nymphs of P. 
dorsata (Say) and P. pictetii is difficult, es- 
pecially immature specimens. The transcon- 



VOLUME 101, NUMBER 2 



329 



tinental species, P. dorsata has been re- 
ported from all surrounding states and Ca- 
nadian provinces except South Dakota 
(Stark et al. 1986). 

Pteronarcys sp. 

Records. — Cass Co., Stearn, 25 Aug 

1960, 1 N (NDSU); Cavalier Co., 20 Oct 
1962, 1 N (NDSU); Grand Forks Co., Tur- 
tle River State Park, 8 Aug 1962, R. D. 
Gordon, 2 N (NDSU); Forest River, Hwy 
18, 3 mi NE Inkster, 1 July 1970, R. Stoaks, 
1 N (NDSU); same but 25 Aug 1970, 1 N 
(NDSU); same but 19 June 1971, 1 N 
(NDSU); same but 28 Aug 1971, 3 N 
(NDSU); same but 15 Sept 1991, 3 N 
(NDSU); Ward Co., Mouse River, Nedrose 
#1, SB Minot, 13 July 1957, R. Nelson, 1 
9 (MSUM). 

The nymphs collected by Ralph D. 
Stoaks are apparently the specimens listed 
by him as P. dorsata, a species that remains 
unconfirmed for the state. The adult female 
listed above could not be determined to spe- 
cies. 

Acroneuria abnormis (Newman) 

Records. — Richland Co., Mirror Pool, 
Sheyenne River, 23 June 1975, P. K. Lago, 
1 S (UMIC). 

This widespread species was expected in 
North Dakota, and is known from all sur- 
rounding states and Canadian provinces 
(Stark et al. 1986, Huntsman et al. 1999). 

Acroneuria lycorias (Newman) 

Records. — Cass Co., Sheyenne River, 5 
mi E Kindred, 28 June 1996, K. Mundal, 2 
c?, 3 $ (NDSU), same but 30 June 1996, 
K. Mundal, 2 c?, 3 $ (NDSU); Grand Forks 
Co., Forest River, Hwy 18, 3 mi NE Inkster, 
16 Oct 1971, R. Stoaks, 1 N (NDSU); Pem- 
bina Co., Pembina River, Walhalla, 9 Sept 

1961, R. L. Post and H. Osborn, 14 N 
(NDSU); same but 20 Oct 1962, E. Saug- 
stad, 2 N (NDSU); same but 20 Oct 1996, 
D. G. Aarhus, 1 N (NDSU). 

This widespread eastern and upper mid- 
western Nearctic species has been recorded 



from adjacent Minnesota and Manitoba. 
Harden and Mickel (1952) indicated that 
this species is one of the few stoneflies that 
occur in the western prairie regions of Min- 
nesota. 

Acroneuria sp. 

Records. — Pembina Co., Pembina River, 
Walhalla, 9 Aug 1961, H. Osborn, 1 N 
(NDSU); Walsh Co., Forest River, USGS 
Gage, 3 mi SE Fordville, 15 Sept 1971, R. 
D. Stoaks, 1 N (NDSU). 

Other possible species of Acroneuria re- 
ported from adjacent states and Canadian 
provinces include A. carolinensis (Banks) 
and A. internata (Walker) (Stark et al. 
1986). Stoaks (1975) Usted A. arenosa, 
however, this nymphal determination is 
doubtful since this species is restricted to 
the eastern U.S. (Stark and Gaufin 1976). 

Perlesta decipiens (Walsh) 

Records. — Cass Co., Sheyenne River, 5 
mi E Kindred, 28 June 1996, K. Mundal, 3 
9 (NDSU); NDAC, Fargo, USDA UV light 
trap, 6 July 1956, 1 9 (NDSU); Richland 
Co., 13 July 1962, 1^,49 (NDSU); Trail 
Co., Elm River, 8 Aug 1969, R. L. Post, 1 
6 (NDSU). 

The occurrence of this geographically 
widespread species was expected in the 
state. 

Perlesta xube Stark and Rhodes 

Records. — Walsh Co., Forest River, Hwy 
35, 8 July 1966, R. L. Post, 2 6,2 9, 3 N 
(NDSU). 

Perlesta xube was recently described 
from a small stream in Cherry County, Ne- 
braska (Stark and Rhodes 1997). This rec- 
ord from the Forest River represents a 
northeastern range extension, indicating 
that this species may occur in remnant 
stream systems throughout the northern 
Great Plains. 

Perlesta sp. 

Records. — Cass Co., Fargo, 13 July 
1973, P K. Lago, 1 9 (UMIC); Dunn Co., 



330 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Knife River, Manning, 29 May 1991, Mott, 
1 N (DSUC); Grand Forks Co., Turtle Riv- 
er, 23 June 1970, D. M. Huntsinger, 1 N 
(NDSU); Forest River, Hwy 18, 3 mi NE 
Inkster, 23 June 1970, R. D. Stoaks, 4 N 
(NDSU); Slope Co., Burning Coal Vein, 
0.25 mi W Logging Camp Ranch, Little 
Missouri National Grassland, 6 July 1968, 
R. D. Stoaks, 6 N (NDSU); Walsh Co., For- 
est River, 3.5 mi W Fordville, Hwy 32, 19 
June 1971, R. D. Stoaks, 31 N (NDSU); 
same but 11 July 1971, R. D. Stoaks, 4 N 
(NDSU); same but at USGS gage, 19 May 
1971, R. D. Stoaks, 1 N (NDSU). 

Nymphs of Perlesta are difficult to spe- 
cifically distinguish, and much of the above 
material is poorly preserved or represents 
early instars. Stoaks (1975) and Neel ( 1985) 
reported P. placida as nymphs and as 
nymphs and adults, respectively. Stark 
(1989) determined that P. placida was a 
complex of at least twelve (now sixteen) 
species. No adults from Neel (1985) were 
available for study. With no associated 
males, the single female adult from Fargo 
was left undetermined. 

Isoperla bilineata (Say) 

Records.— Cass Co., Fargo, NDSU Cam- 
pus, 7 June 1963, R. W. Poole, 7 d, 1 9 
(NDSU); same but 16 June 1963, R. W. 
Poole, 7 d, 1 9 (NDSU) same but 18 June 
1993, 6 6 (NDSU); Sheyenne River, 5 mi 
E Kindred, 30 June 1996, K. Mundal, 1 d, 
4 9 (NDSU); NDAC, Fargo, USDA UV 
light trap, 9 June 1956, 20 (5, 3 9 (NDSU). 
Emmons Co., 16 mi W Linton, 20 June 
1975, P K. Lago, 2 6 (UMIC); Grant Co., 
Heart Butte Dam, 19 June 1975, P K. and 
B. A. Lago, 2 S (UMIC); Pembina Co., 
Tongue River, Kotchman Farm, near Cav- 
alier, 29 June 1974, R K. and B. A. Lago, 
3 S (UMIC); same but 2 July 1975, 2 6 
(UMIC). 

Stoaks (1975) previously reported this 
species, but no specimens from his study 
were available for examination. Szczytko 
and Stewart (1978) indicated material from 



the Red River of the North, Minnesota, 
which forms the border with North Dakota. 

Isoperla longiseta (Banks) 

Records. — Billings Co., Little Missouri 
River, Elkhom Ranch, 20 June 1965, R. L. 
Post, 1 9 (NDSU); Cass Co., Fargo, 11 
May 1939, D. G. Denning, 1 9 (UMSP); 
Slope Co., Little Missouri River, at Mar- 
marth, Hwy 12, 14 July 1997, R. W. Bau- 
mann and B. C. Kondratieff, 6 d, 7 9 
(BYUC, CSUC). 

Isoperla longiseta is known from west- 
ern and midwestern North America, and oc- 
curs further east than any other typically 
western Isoperla (Szczytko and Stewart 
1979). It is often considered a typical prai- 
rie stonefly of larger streams and rivers 
(Harden and Mickel 1952, Ricker 1946, 
1964). Harden and Mickel (1952) previous- 
ly mentioned the 1939 record of/, longiseta 
from Fargo. 

Haploperla orpha (Frison) 

Records. — Pembina Co., Tongue River, 
Kotchman Farm near Cavalier, 21 June 
1974, R K. and B. A. Lago, 2 d, 3 9 
(BPSC); same but 23 July 1975, 1 c?, 3 9 
(UMIC). 

Haploperla orpha is a relatively poorly 
known species having also been recorded 
from Maine, Minnesota, Quebec, New 
Brunswick, and Wisconsin (Stark et al. 
1986). Typically, nymphs of this species are 
found in medium and small-sized streams 
with some gradient. 

Acknowledgments 

We thank Dr. B. P. Stark for reviewing 
the manuscript, providing the SEM photo- 
graphs and material. Drs. P. J. Clausen, Uni- 
versity of Minnesota; R. D. Gordon, Wil- 
low City, North Dakota; P. K. Lago, Uni- 
versity of Mississippi; C. P. Milne, Jr., 
Dickinson State University; D. A. Rider, 
North Dakota State University; R. A. Roy- 
er, Minot State University; and P. D. Per- 
kins, Museum of Comparative Zoology, 



VOLUME 101. NUMBER 2 



331 



Harvard University, are thanked for loaning 
valuable material. 

Special thanks are given to Charles P. 
Milne, Jr. for making a concerted effort to 
collect winter stonefly species, which re- 
sulted in the discovery of Oemopteryx fos- 
ketti. Dr. Gerald M. Fauske, NDSU, helped 
us locate questionable localities from across 
the state. 

Literature Cited 

Baumann. R. W., A. R. Gaufin. and R. F. Surdick. 
1977. The stoneflies (Plecoptera) of the Rocky 
Mountains. Memoirs of the American Entomolog- 
ical Society 3 1 . 208 pp. 

DeWalt, R. E., B. R Stark, and M. A. Harris. 1998. 
Perlesta golconda (Plecoptera: Perlidae), a new 
stonefly species from Illinois. Entomological 
News 109: 315-317. 

Gordon, R. D. and R. L. Post. 1965. North Dakota 
water beetles. North Dakota Insects. Publication 
5. Agricultural Experiment Station, North Dakota 
State University 53 pp. 

Harden, R H. and C. E. Mickel. 1952. The stoneflies 
of Minnesota (Plecoptera). University of Minne- 
sota Agricultural Experiment Station, Technical 
Bulletin 201. 82 pp. 

Huntsman, B. O., R. W. Baumann. and B. C. Kondra- 
tieff. 1999. The stoneflies (Plecoptera) of the 
Black Hills of South Dakota and Wyoming, USA: 
Distribution and zoogeographic affinities. Great 
Basin Naturalist 59: 1-17. 

Kirchner, R. E and B. C. Kondratieff. 1997. A new 
species of Nearctic Perlesta (Plecoptera: Perlidae) 
from Virginia. Proceedings of the Entomological 
Society of Washington 99: 290-293. 

Neel, J. K., Sr. 1985. A northern prairie stream. Uni- 
versity of North Dakota Press, Grand Forks. 274 
pp. 

Poulton. B. C. and K. W. Stewart. 1991. The stoneflies 
of the Ozark and Ouachita Mountains (Plecop- 
tera). Memoirs of the American Entomological 
Society 38, 1 16 pp. 

Ricker, W E. 1946. Some prairie stoneflies (Plecop- 



tera). Transactions of the Royal Canadian Institute 
26: 3-8. 

. 1964. Distribution of Canadian stoneflies, pp. 

50-71. In lilies, J. ed., 3. Internationales Sympo- 
sium iiber Plecopteren. August. Gewasser und Ab- 
wasser. Bagel, Dusseldorf. 138 pp. 

Ricker, W. E. and H. H. Ross. 1968. North American 
species of Taeniopteryx (Plecoptera: Insecta). 
Journal of the Fisheries Research Board of Canada 
25: 1423-1439. 

Ross. H. H. and W. E. Ricker. 1971. The classification, 
evolution, and dispersal of the winter stonefly ge- 
nu?, Allocapnia. Illinois Biological Monograph 45, 
166 pp. 

Stark. B. P. 1989. Perlesta placida (Hagen), an eastern 
Nearctic species complex (Plecoptera: Perlidae). 
Entomologica Scandinavica 20: 263-286. 

Stark, B. P and A. R. Gaufin. 1976. The Nearctic spe- 
cies of Acroneiiria (Plecoptera: Perlidae). Journal 
of the Kansas Entomological Society 49: 221- 
253. 

Stark, B. P and H. A. Rhodes. 1997. Perlesta xube, a 
new stonefly species from Nebraska (Plecoptera: 
Perlidae). Entomological News 108: 92-96. 

Stark, B. R, S. W. Szczytko, and R. W. Baumann. 
1986. North American stoneflies (Plecoptera): sys- 
tematics, distribution, and taxonomic references. 
Great Basin Naturalist 46: 383-397. 

Stewart, K. W. and B. P Stark. 1988. Nymphs of North 
American stonefly genera (Plecoptera). Thomas 
Say Foundation, Entomological Society of Amer- 
ica. 12, 460 pp. 

Stoaks, R. D. 1975. Seasonal and spatial distribution 
of riffle dwelling aquatic insects in the Forest Riv- 
er, North Dakota. Ph.D. dissertation. North Dakota 
State University, Fargo. 163 pp. 

Szczytko, S. W. and K. W. Stewart. 1978. Isoperla 
bilineala: designation of a neotype and allotype, 
and further descriptions of egg and nymph. An- 
nals of the Entomological Society of America 71: 
212-217. 

. 1979. The genus Isoperla (Plecoptera) of 

western North America; holomorphology and sys- 
tematics, and a new stonefly genus Cascadoperla. 
Memoirs of the American Entomological Society. 
32: 120 pp. 

Van der Valk, A. 1989. Northern Prairie Wetlands. 
Iowa State University Press, Ames. 400 pp. 



PROC. ENTOMOL. SOC. WASH. 

101(2). 1999, pp. 332-336 

A NEW NEARCTIC SPECIES OF LESTODIPLOSIS 
(DIPTERA: CECIDOMYIIDAE) PREYING ON AN OAK LEAF TIER, 
PSILOCORSIS QUERCICELLA (LEPIDOPTERA: OECOPHORIDAE) 

Raymond J. Gagne and John T. Lill 

(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- 
tution, Washington, DC 20560-0168, U.S.A. (e-mail: rgagne@sel.barc.usda.gov); (JTL) 
Department of Biology, University of Missouri-St. Louis, St. Louis, MO 63121, U.S.A. 



Abstract. — A new species of cecidomyiid, Lestodiplosis venusta Gagne, is described, 
illustrated, and compared to its most similar congeners. Larvae of the new species were 
found in Missouri preying on caterpillars of Psilocorsis quercicella Clemens, a leaf skel- 
etonizer of oaks. An unidentified species of Ceraphronidae was reared from pupae of the 
Lestodiplosis. 

Key Words: Lestodiplosis, predator, oak leaf tier 



A new species of Lestodiplosis attacking 
larvae of the oecophorid Psilocorsis quer- 
cicella Clemens was discovered by one of 
us (JTL) during the course of an ecological 
study of the natural enemies of the lepidop- 
teran in Missouri. Psilocorsis quercicella is 
a common bivoltine moth found throughout 
eastern United States (Covell 1984). The 
larvae skeletonize the surface of oak leaves 
within leaf shelters formed by tying togeth- 
er adjacent oak leaves with silk. Larvae of 
the gall midge predator appear to be highly 
mobile ectoparasites of early instar P. quer- 
cicella larvae and are often gregarious. 
When present, one to four of the preda- 
ceous cecidomyiids were found per cater- 
pillar. The predator was found feeding on 
P. quercicella on both white oak (Quercus 
alba L.) and black oak (Q. velutina Lam.). 
After feeding, full grown Lestodiplosis lar- 
vae spin a silken cocoon within the leaf 
shelter, often incoiporating pieces of cater- 
pillar frass onto the surface of the cocoons. 
To date, larvae of the predator have been 
observed only from second generation P. 



quercicella larvae collected in late summer 
and fall in Missouri. An unidentified gre- 
garious parasitoid (Hymenoptera: Cera- 
phonidae) has been reared from the Lesto- 
diplosis pupae. 

The genus Lestodiplosis contains some 
175 known species in the world that attack 
many kinds of insects and mites. Some ap- 
pear to be specialist predators, others are 
evidently generalists. Many species are 
known from only one or a few specimens 
so it is especially helpful to have several 
series of the same species from a particular 
niche, such as JTL found in leaf ties of P. 
quercicella. 

Methods 

Larvae of the predator were reared to the 
adult stage in 16-oz. closed clear plastic 
containers kept at 23 °C and with a photo- 
period of 14:10 (L:D) h. Humidity was 
maintained with moist filter paper. Speci- 
mens of immature stages and reared adults 
were preserved in 70% isopropyl alcohol. 
Specimens were mounted on microscope 



VOLUME 101, NUMBER 2 



333 



slides using the method outhned in Gagne 
(1989). Terminology for adult morphology 
follows usage in McAlpine et al. (1981) and 
for larval morphology that in Gagne ( 1989). 

Lestodiplosis venusta Gagne, 
new species 

(Figs. 1-13) 

Adult. — Head: Eyes connate, 11-12 fac- 
ets long at vertex; facets hexagonal, all 
closely adjacent. Occiput with dorsal pro- 
tuberance. Frons with 10—14 setae. Labella 
hemispherical but pointed apically, each 
with several lateral setae. Palpus 4-seg- 
mented. Male antennal flagellomeres (Fig. 
1) binodal; basal node and the distal third 
of the neck dark; one circumfilum on the 
basal node, two on the distal, the loops of 
the three circumfila unequal in length; the 
proximal and distal circumfila with ventral- 
most loops greatly elongated and much lon- 
ger than remaining loops. Female flagel- 
lomeres (Fig. 2) with basal part of node and 
distal part of neck darker than remainder of 
flagellomere; circumfila with some short 
loops. 

Thorax: Wing (Fig. 3) 1.8-2.3 mm long, 
R5 slightly curved, joining C posterior to 
wing apex; 7-8 irregular dark spots present 
on wing. Legs (Fig. 4) with alternating light 
and dark groups of scales. Tarsal claws 
(Fig. 5) untoothed, curved beyond mid- 
length; empodia attaining bend in claws. 

Male abdomen: First through seventh 
tergites entire, rectangular, with single pos- 
terior row of setae, several lateral setae, 
scattered scales, and 2 anterior trichoid sen- 
silla; eighth tergite undifferentiated, the 
only vestiture the anterior pair of trichoid 
sensilla. First through seventh sternites 
quadrate, with single to double posterior 
row of setae, scattered setae elsewhere, no 
scales, and 2 anterior trichoid sensilla; 
eighth sternite similar to preceding except 
weakly sclerotized anterolaterally. Genitalia 
(Figs. 6-8): cerci each with rounded pos- 
terior margin; hypoproct simple, rounded at 
apex, as long as but slightly wider in ventral 
view than cerci, with 2 pairs of apicoventral 



setae, produced anteroventrally into large 
lobe lying in close juxtaposition with dorsal 
surface of aedeagus and covered posteriorly 
with short spinules; aedeagus elongate with 
blunt apex, sinuous in side view; gonocox- 
ite elongate cylindrical with short, triangu- 
lar mesobasal lobe bearing short spinules; 
gonostylus elongate cylindrical, with setu- 
lae near base and covered with minute ca- 
rinae and only several short setae beyond. 

Female abdomen (Figs. 9-11): First 
through seventh tergites entire, rectangular, 
with mostly single row of posterior setae, 
several lateral setae, extensively covered 
with scales, and 2 anterior trichoid sensilla. 
Second through seventh sternites quadrate, 
extensively covered with setae and scales 
and with anterior pair of trichoid sensilla. 
Eighth segment much narrowed, without 
differentiated tergite and sternite, the ter- 
gum with anterior pair of trichoid sensilla 
the only vestiture, the sternum with anterior 
pair of trichoid sensilla and covered with 
setae some distance beyond. Tenth segment 
without vestiture dorsally, with scattered se- 
tae ventrally; cercus large, ovoid, with ven- 
tral field of short, closely-set, sensory setae, 
and scattered setae elsewhere. Hypoproct 
short, wide, the convex posterior edge with 
2 short setae. Apex of ninth sternite pro- 
truding posteriorly below hypoproct, rigid, 
triangular. 

Third larval instar (Figs. 12-14). — Integ- 
ument mostly smooth, 2 pseudopods pres- 
ent ventrally on mesothorax and metathorax 
and 3 each on first through seventh abdom- 
inal segments. Antenna much longer than 
wide. Cephalic apodemes slightly longer 
than head capsule. Spatula absent. Lateral 
papillae in 2 groups of 3 on each side of 
central line, only one setose in each group. 
Second and fifth dorsal papillae of each 
segment much shorter than remaining four. 
Terminal segment with 6 elongate setae, 
usually with expanded apices. 

Holotype. — 6 , reared from larva preying 
on larva of Psilocorsis quercicella on oak, 
Cuivre River State Park, Troy, Missouri, 1- 
IX- 1997, J. Lill, deposited in the National 



334 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 







^^ 





8 



Figs. 1-8. Lestodiplosis vemista. L Male third antennal flagellomere. 2, Female third antennal flagellomere. 
3, Wing. 4, Foreleg. 5, Tarsal claw and empodium. 6. Gonopod. cerci, hypoproct, and aedeagus (dorsal). 7, 
Cercus, hypoproct, and aedeagus (lateral). 8, Hypoproct (ventral). 



VOLUME 101. NUMBER 2 



335 




10 




13 



14 



Figs. 9-14. Lestodiphsis venmta. 9-1 1. Female postabdomen. 9. Seventh tergite to cerci (not all setal bases and 
setae indicated; dorsal). 10, Cerci, hypoproct and apex of ninth sternum, the postabdomen retracted (lateral). 11, 
Seventh stemite to cerci (not all setal bases and setae indicated; ventral). 12-14. Larva. 12. Head. neck, and first two 
thoracic segments (dorsal). 13, part of mesothorax with two groups of three lateral papillae, a ventral papilla, and two 
pseudopods (ventrolateral). 14. Apex of seventh and all of eighth and terminal abdominal segments (dorsal). 



336 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



Museum of Natural History (USNM), 
Washington, DC. 

Other material examined. — All associat- 
ed with leaf ties of Psilocorsis quercicella, 
Cuivre River State Park, Troy, Missouri: 6 
(?, 5 9,9 larvae, same data as holotype; 6 
S, \ 9, same data as holotype except col- 
lected ll-IX-1996, all in USNM. 

Etymology. — The specific name, venus- 
ta, is an adjective meaning beautiful, with 
reference to the striking black and white 
banded legs and antenna. 

Remarks. — The banded antenna and legs 
and the spotted wings of this species are 
generally similar to some other Lestodi- 
plosis species, but the combination of un- 
equal male circumfila and a blunt-tipped ae- 
deagus is characteristic in the Nearctic Re- 
gion of a narrower group of three species 
that differ chiefly in details of the male gen- 
italia. These three species are: Lestodiplosis 
florida Felt, Lestodiplosis cinctipes (Felt), 
and Lestodiplosis satiata Felt. These other 
species are now under investigation by one 
of us (RJG) for a revision in progress of 
Nearctic Lestodiplosis. 

Compared to the new species, the ventral 
hypoproctal lobe of L. florida is covered by 
fewer but more robust spinules and its ae- 
deagus is much wider on the basal two- 
thirds and tapers abruptly to a narrow neck. 
Lestodiplosis florida was originally taken 
from a leaf roll on a Crataegus in Florida. 
This species is represented in the USNM 
collection by a specimen from Florida that 
was reared from galls on Croton linearis 
Jacq. and another that was swept from Tsu- 
ga canadensis (L.) Carr. in Ontario. 

The ventral lobe of the hypoproct of both 
L. cinctipes and L. satiata is covered more 
extensively and by finer spinules than in the 
new species and the aedeagus in these spe- 
cies is narrower throughout its length than 
on L. venusta. In addition, the antennal cir- 
cumfila loops of both species are all much 
longer than in L. venusta. Both species were 
originally caught in flight in New York, but 
a specimen of L. satiata in the USNM was 



reared in association with cynipine leaf 
galls on Quercus sp. in Pennsylvania and 
another was reared from flower heads of 
Borrichia frutescens (L.) DC. in North Car- 
olina. 

There is one other Nearctic Lestodiplosis 
species, besides L. venusta, that has been 
recorded as feeding on caterpillars. Felt 
(1933) described Lestodiplosis novangliae 
Felt from specimens associated with a tor- 
tricid Epinotia "nanana'' on spruce in 
Massachusetts. He noted also that larvae 
similar to that species were found on an- 
other tortricid, Rhyacionia sp. on pine. The 
gonocoxites of L. novangliae, unlike those 
of L. venusta, have prominent and spiny 
mesobasal lobes. 

Acknowledgments 

We thank B. Schuette and the Missouri 
Department of Natural Resources for logis- 
tical support and collection permits, P. Ma- 
likul for making the slide preparations, C. 
Anderson for computer assistance in mak- 
ing the plates, and, for their comments on 
drafts of the manuscript: P. Kolesik, Uni- 
versity of Adelaide, South Australia; A. L. 
Norrbom of the Systematic Entomology 
Laboratory, USDA, Washington, D.C.; M. 
Solinas, Universita degli Studi di Perugia, 
Italy; and J. Yukawa, Kyushu University, 
Fukuoka, Japan. The field research was 
funded with a National Science Foundation 
Dissertation Improvement Award (DEB- 
9700887) to JTL. 

Literature Cited 

Covell, C. v.. Jr. 1984. A Field Guide to the Moths of 
Eastern North America. Houghton Mifflin Co., 
Boston. 

Felt. E. R 1933. A new Lestodiplosis. Psyche 40: 1 13- 
114. 

Gagne. R. J. 1989. The Plant-Feeding Gall Midges of 
North America. Cornell University Press, Ithaca, 
New York, xiii + 355 pp. & 4 pis. 

McAlpine, J. F, B. V. Peterson, G. E. Shewell, H. J. 
Teskey, J. R. Vockeroth, and D. M. Wood, Eds. 
1981. Manual of Nearctic Diptera, Vol. I, Re- 
search Branch, Agriculture Canada, Monograph 
27, vi -I- 674 pp. 



PROC. ENTOMOL. SOC. WASH. 

101(2). 1999. pp. 337-346 

DESCRIPTION OF TWO NEW SPECIES OF AMBLYCERUS THUNBERG 

(COLEOPTERA: BRUCHIDAE) WITH A PROBABLE 

STRIDULATORY MECHANISM 

ClBELE S. RlBEIRO-COSTA 

Departamento de Zoologia, Universidade Federal do Parana, C. P. 19020, 81.531-990, 
Curitiba, Parana, Brazil (e-mail: stra@bio.ufpr.br) 



Abstract. — Amblycerus atypicus, new species, and A. ischiodontus, new species, are 

described and illustrated. The former has an area with overlapping scales in part of the 
external elytral margins and the inner distal portion of the hind femur with fine transverse 
striation. Amblycerus ischiodontus has an area transversely striate on the metepisternum 
and an apical tooth on the ventral margin of the hind femur. The areas on the elytra and 
metepisternum probably act as a file and the striate area or the tooth of the hind femur 
as the scraper The elytro-femoral method of stridulation is postulated for the first time 
in Bruchidae and is shared with Amblycerus cistelimis (Gyllenhal), A. jatayensis (Pic), A. 
sosia Ribeiro-Costa and Kingsolver, A. whiteheadi Kingsolver and A. guazumicola King- 
solver and Johnson. The other kind, involving the metepisternum and hind femur, has 
already been described for three other Amblycerus species: A. eustrophoides (Schaeffer), 
A. pollens (Sharp), and A. stridulator Kingsolver, Romero N., and Johnson. A key for the 
bruchid species with modified body areas probably involved in stridulation is presented. 

Key Words: Amblycerus, stridulation, taxonomy, Bruchidae 



Kingsolver (1970) first recorded the pres- 
ence of areas of the integument probably 
involved in stridulation in Bruchidae. He 
indicated the presence of a fusiform node 
with transverse striations on the metepister- 
num and the presence of an apical blunt 
tooth on the ventral margin of the hind fe- 
mur for Amblycerus eustrophoides (Shaef- 
fer), which has a distribution restricted to 
North America. Kingsolver et al. ( 1993) no- 
ticed the same stridulatory areas for A. stri- 
dulator described from Mexico, Costa Rica, 
and Venezuela and for A. pollens (Sharp) 
recorded from Belize, Costa Rica, and Bra- 
zil. 

Amblycerus atypicus, n. sp. (French Gui- 
ana), has an area with overlapping scales on 
the external margins of the elytra and fine 
transverse striation on the inner distal por- 



tion of the hind femur The other new spe- 
cies, Amblycerus ischiodontus (Brazil), has 
the metepisternum and hind femur with ar- 
eas transversely striate, similar to those 
mentioned for A. eustrophoides, A. pollens, 
and A. stridulator. These areas for each spe- 
cies are in contact when the hind leg is 
moved, and, even though no sound emis- 
sion has been perceptible from dead speci- 
mens, it is possible that they form a strid- 
ulatory mechanism. Since males and fe- 
males of both species have these areas, I 
hypothesize that the mechanism probably is 
not related with courtship behavior 

The type of stridulation in A. atypicus 
can be characterized as the elytro-femoral 
method, established by Dumortier (1963). 
The area with overlapping scales on the el- 
ytron would be the "pars stridens" (file) 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



and the striate area of the hind femur, the 
"plectrum" (scraper). Dumortier (1963) 
cited the presence of the elytro-femoral 
method for the Coleopteran families Scar- 
abaeidae, Lucanidae, Cerambycidae, Cicin- 
delidae, Tenebrionidae, and Carabidae. This 
method of stridulation is hypothesized for 
the first time in Bruchidae and is shared 
with the following Amblycerus Thunberg 
species: A. cistelinus (Gyllenhal), A. jatay- 
ensis (Pic), A. sosia Ribeiro-Costa and 
Kingsolver, A. whiteheadi Kingsolver, and 
A. guaziimicola Kingsolver and Johnson. 

The methodology used for the descrip- 
tions follows Ribeiro-Costa (1997). 

Amblycerus atypicus Ribeiro-Costa, 
new species 

(Figs. 1-10) 

Description. — Measurements (pronotitm 
+ elytra): Length 8.17-8.83 mm; width 
4.83-5.17 mm. 

Integument: Antenna dark red. Dorsal 
surface (Fig. 1) dark red with scattered 
black spots on elytra, two on basal one- 
third, four at middle one-third and two on 
apical one-third. Lateral margins of elytra 
gently darker. Undersurface and legs dark 
red. Pygidium with large, ovate, central, 
dark brown spot (Fig. 4). 

Vestiture: General coloration on dorsal 
surface golden yellow (Fig. 1). Pronotum 
with white setae forming six small, round 
spots arranged in triangle in middle lateral 
and scattered patches along basal lobe (Fig. 
1). Scutellum usually white. Elytra (Fig. 1) 
with white setae in scattered patches along 
basal margin and around scutellum, often in 
a few small, irregular spots on median and 
external margins, in lines along interstitial 
margins and strial sulci, and condensed into 
a few, very small irregular patches on re- 
mainder of elytra. Pygidium (Fig. 4) with 
large, ovate, velvety dark brown spot, bor- 
dered with dense white setae and with two 
small patches of dark brown pillosity in an- 
terolateral areas. Undersurface gently mot- 
tled brown and white. Lateral margins of 



abdominal sterna with white setae gently 
condensed into round patches. 

Frons slightly flat, evenly punctate ex- 
cept on frontal carina. Eyes moderately fac- 
eted (3 ommatidia), protruding laterally; oc- 
ular index 3.9; ocular sinus 1/3.8 diameter 
of eye in lateral view; postocular lobe 1/9.5 
times largest length of eye in lateral view. 
Antenna (Fig. 5) serrate from fourth to 
tenth segments, all gently longer than wide 
(1.1 times); last segment elliptical. Prono- 
tum trapezoidal with lateral margins slight- 
ly arched in dorsal view, cervical sulcus 
present in lateral third, a sulcus outlining 
the emargination of basal lobe; long lateral 
carina, nearly reaching apex of pronotum, 
not forked anteriorly; surface densely punc- 
tulate, lateral one-third on either side also 
coarsely punctate. Prostemal process wide, 
margins sulcate, apex subacute, moderately 
expanded exceeding fore coxae and fitting 
into sulcate mesosternum. Scutellum (Fig. 
2) about 1.9 times as long as wide, with 
tridentate apex. Elytra slightly convex in 
cross section along elytral suture, with an 
area with overlapping scales in part of the 
external margins (Fig. 9), truncated apically 
and with striae strongly impressed. Mete- 
pisternum with some punctures moderately, 
sparsely coarser; transverse axis of mete- 
pistemal sulcus (Fig. 3) strongly divergent 
from metapleural suture, very short longi- 
tudinal axis, less than half metepisternal 
length, not modified in a fusiform node 
with transverse striations (Fig. 3). Hind 
coxa with punctures moderately coarse and 
slightly dispersed, except in basal third. 
Hind femur (Fig. 6) 2.8 times as long as 
wide; apical tooth on ventral margin absent; 
internal face on distal portion with trans- 
verse striations (Figs. 6, 10). Hind tibia 
(Fig. 6) with coronal teeth approximately of 
same size; lateral spur 2.9 times length of 
median, first hind tarsal segment about 1.2 
times as long as lateral spur and 3.6 times 
median spur; ventral face slightly convex 
with inconspicuous lines of punctures and 
setae in margins. Male pygidium vertical in 
lateral view, female oblique; male eighth 



VOLUME 101, NUMBER 2 



339 




^, J Dark red integumait 
siisl Golden yellow hairs 

I Black integument 
Golden yellow hairs 

I Dark red integument 
I White hairs 

I Dark brown integument 
I Dark brown hairs 





1 mm 



Figs. 1-6. Amblycenis atypicus. 1, Dorsal habitus. 2, Scutellum. 3, Lateral view of head and thorax 4 
Pygidium. 3, Antenna. 6. Hind leg. 



340 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





Figs. 7-8. Ambhcents atypicus. 7, Male terminalia. median lobe. 8, Same, tegmen and lateral lobes. 



tergite acute. Fifth visible abdominal ster- 
num of female longer than that of male at 
middle, margin in both sexes entire. 

Male terminalia (Figs. 7, 8): Eighth ter- 
gite acute apically. Median lobe (Fig. 7) 
with length 4.2 times its largest width in 



basal area; ventral valve moderately long, 
acute apically, with straight lateral margins; 
dorsal valve rounded. Basal area of internal 
sac (Fig. 7) without anterior and median 
sclerites; a pair of posterior tooth-like scler- 
ites. Median region of internal sac (Fig. 7) 



VOLUME 101, NUMBER 2 



341 



with two central, long laminar sclerites each 
with angulate apical portion and each one 
with one or two teeth near base, one sub- 
basal and two on median portion; pair of 
sclerites laterad to laminars, longer than 
wide, each with one small curved tooth di- 
rected upward; unpaired median sclerite, 
0.3 times as long as laminars, slightly sin- 
uate in lateral view, with one side serrate. 
In apical area of internal sac (Fig. 7) one 
sclerite with long stems among many den- 
ticles. Tegmen with shallow emargination 
among enlarged lateral lobes (Fig. 8). 

Discussion. — Romero et al. ( 1996) estab- 
lished the cistelinus group including A. cis- 
telimis, A. sosia, and A. giiazumicola main- 
ly considering both the integument and pu- 
bescence pattern on the dorsal surface, not 
mentioning the presence of areas on elytra 
or hind femur possibly involved in stridu- 
lation. Amblyceriis atypicus (French Gui- 
ana), A. jatayensis (Brazil) and A. white- 
headi (Panama, Nicaragua, Costa Rica, Co- 
lombia), probably will be included in this 
group (revisionary study currently under- 
way) because they share characters cited by 
Romero et al. (1996) and also the areas on 
the external margins of the elytra with over- 
lapping scales (Fig. 9) and transverse stri- 
ation on the inner distal portion of hind fe- 
mur (Fig. 10), which are apparently strid- 
ulatory in function. 

Amblyceriis atypicus is clearly separated 
from all the other Amblyceriis species and 
especially from those in the cistelinus group 
by the form, number and arrangement of 
sclerites in the male internal sac (Fig. 7). 

Types.— Holotype 9. FRENCH GUI- 
ANA: Maroni River; Collection Wm 
Schaus, deposited in the National Museum 
Natural of History, Smithsonian Institution, 
Washington, D.C. U.S.A. One paratype 6 
with same label deposited in the Colegao 
de Entomologia Pe. Jesus S. Moure, Curi- 
tiba, Brazil. 

Etymology. — The species name refers to 
the atypical elytro-femoral method of pos- 
sible stridulation in Amblyceriis. 



Amblycerus ischiodontus Ribeiro-Costa, 
new species 

(Figs. 11-18) 

Description. — Measurements (pronotum 
+ elytra): Length 4.48 mm; width 2.72 
mm. 

Integument: General coloration, dark 
red. Eyes, legs and undersurface of thorax 
darker. 

Vestiture: Head, appendages and ventral 
area of thorax and abdomen, flavous; met- 
epistemum in part, golden yellow. Prono- 
tum (Fig. 11) golden yellow rarely with 
small brown patches. Scutellum flavous. El- 
ytra (Fig. 1 1 ) golden yellow; brown hairs 
in rare irregular patches and flavous hairs 
in lines along interstitial margins and strial 
sulci. Pygidium (Fig. 11) golden yellow 
with two small central, brown patches. 

Frons slightly flat, evenly punctuated, 
frontal carina absent. Eyes moderately fac- 
eted (7 ommatidia), protruding laterally; oc- 
ular index 3.8; ocular sinus 1/5.0 diameter 
of eye in lateral view; postocular lobe 1/6.7 
times largest length of eye in lateral view. 
Antenna (Fig. 16) serrate from fourth to 
tenth segments, all perceptibly longer than 
wide (1.4 times), last segment subelliptical. 
Pronotum trapezoidal with lateral margins 
moderately arched in dorsal view, cervical 
sulcus in lateral thirds and a sulcus outlin- 
ing basal lobe; long lateral carina, nearly 
reaching apex of pronotum, not forked an- 
teriorly; surface densely punctulate, lateral 
one-third either side also coarsely punctate. 
Prostemal process wide, not sulcate, apex 
subacute, moderately expanded between an- 
terior coxae and fitting into sulcate meso- 
sternum. Scutellum (Fig. 12) about 1.8 
times as long as wide, apex strongly triden- 
tate on rounded base. Elytra slightly convex 
in cross section along elytral suture, sub- 
truncated apically and with striae strongly 
impressed. Transverse axis of metepister- 
num (Fig. 13) moderately divergent from 
metapleural suture; transversely striate and 
strongly curved area running with longitu- 
dinal axis, apparently being a modification 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 9-10. Amhlycenis atypicus. 9, Scanning electron micrograph (SEM) of the external margin of elytron 
showing "tile." 10, SEM of the inner distal portion of the hind femur showing "scraper." 



VOLUME 101, NUMBER 



343 




'iV'i' 
".'."ii 

».Vi".'; 



Dark red integument 
Golden yellow hairs 

Dark red integument 
I Brown hairs 

□ Dark red integument 
Flavous hairs 



1 mm 




Figs. 11-16. Amblycems ischiodontiis. 11, Dorsal habitus. 12, Scutellum. 13, Metepisternum. 14, Hind leg. 
15, Ventral margin of hind femur showing the tooth. 16, Antenna. 



of it (Fig. 13). Hind coxa with coarse punc- distal region without transverse striations. 

tures slightly denser in distal half. Hind fe- Hind tibia (Fig. 14) with coronal denticles 

mur (Fig. 14) 2.3 times as long as wide and approximately of same size; lateral spur 2.4 

with tooth pronounced in distal portion of times length of median, first hind tarsal seg- 

ventral margin (Fig. 15); internal face on ment 1.2 times as long as lateral spur and 



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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 




Figs. 17-18. Amblycenis ischiodoiitiis. 17, Male terminalia, median lobe. 18, Same, tegmen and lateral lobes. 



2.8 times median spur; ventral face mod- 
erately convex with inconspicuous lines of 
punctures and setae in margins. Male py- 
gidium vertical, round apically. Fifth visible 
abdominal sternum not emarginate in male. 
Female unknown. 

Male tenninalia (Figs. 17, 18): Eighth 
tergite rounded apically. Median lobe (Fig. 
17) with length 3.8 times its largest width 



in basal area; ventral valve moderately 
long, acute apically, with concave lateral 
margins; dorsal valve subtriangular with 
straight lateral margins and rounded apex. 
Basal area (Fig. 17) of internal sac without 
anterior and median sclerites; a pair of pos- 
terior thin sclerites, prolonged and with a 
tooth sub-basally on one side. Median area 
of internal sac (Fig. 17) with a pair of lam- 



VOLUME 101. NUMBER 2 



345 



inar sclerites, sinuous, serrate sub-basally, 
teeth approximate; unpaired sclerite in fork 
form 0.6 times as long as laminars, with 
moderately separate stems. In apical area of 
internal sac (Fig. 17) one sclerite with very 
long stems. Median and apical areas of in- 
ternal sac with denticles near apical sclerite. 
Tegmen (Fig. 18) with moderate emargi- 
nation in "V" between enlarged lateral 
lobes. 

Discussion. — Amblycerus ischiodontus 
shares with A. eustrophoides, A. pollens, 
and A. sthdidator an area of the metepister- 
num with transverse striations (Fig. 13) and 
an apical tooth on the ventral margin of the 
metafemur (Fig. 15). The study of all char- 
acters together did not show great affinity 
among the mentioned species. Differences 
are easily recognized in the patterns of pu- 
bescence, form of the striate area, and the 
sclerites in the male internal sac. 

Kingsolver et al. (1993) commented that 
A. pollens is more similar to A. eustrophoi- 
des when compared with A. strididator. 
Romero et al. (1996) included A. eustro- 
phoides in the alternatus group along with 
A. serieguttatus, A. alternatus, and A. 
schwarzi and stablished the stridulator 
group only for A. stridulator. 

Revisionary studies on Brazilian species 
under way indicate that A. pollens and A. 
ischiodontus are members of two distinct 
species groups, respectively. 

According to Arrow (1904) the body area 
where the stridulatory mechanism is placed 
in Coleoptera can be the same in non-relat- 
ed groups, as it can be different in different 
members of the same group. According to 
him, there are Coleoptera with the two 
types of stridulatory mechanism placed in 
different parts of the body. 

Amblycerus ischiodontus is easily sepa- 
rated from all other Amblycerus by its char- 
acteristic form and disposition of sclerites 
in the male internal sac (Fig. 17). 

Type. — Holotype 6. BRAZIL: Amazo- 
nas, Parana do Xiboreninho, 03°15'S- 
06°00'W; 5 Aug. 79; mixed water; Canopy 
Fogging Project TRS#05, Tray 326 Adis, 



Erwin, Montgomery et al. collectors; de- 
posited in the National Museum Natural of 
History, Smithsonian Institution, Washing- 
ton, D.C., U.S.A. 

Etymology. — The name of the species 
refers to the presence of a tooth on the ven- 
tral margin of the hind femur. 

Key to Amblycerus Species with 

Areas of Integument Probably 

Involved in Stridulation 

1 . External margins of elytra partly with an area 
with overlapping scales (Figs. 3, 9); inner distal 
portion of hind femur with fine transverse stri- 
ation (Figs. 6, 10); usually scattered black spots 
on elytra (Fig. 1); pygidium with large, ovate, 
velvety dark brown spot (Fig. 4) 2 

- Elytra and hind femur without integumental 
modifications; patterns of coloration on elytra 
and pygidium otherwise 7 

2. Elytra with white hairs vaguely mottled; very 
coarse punctures on lateral third of pronotum; 
male terminalia with four pairs of sclerites and 
one unpaired sclerite 

Amblycerus giiazmnicola (Kingsolver and 

Johnson) 

- Elytra usually with well marked white inegular 
spots on median and external margins and 
small irregular patches on remainder of elytra 
(Fig. 1 ); moderately coarse punctures on lateral 
third of pronotum; number of sclerites in in- 
ternal sac of male terminalia otherwise 3 

3. Male internal sac with three pairs of sclerites 
and one unpaired sclerite (Fig. 7) 

Amblycerus utypicus, new species 

- Male internal sac with two pairs of sclerites 
and one unpaired sclerite 4 

4. Pair of laminar sclerites in male internal sac 
with tooth 5 

- Pair of laminar sclerites in male internal sac 
smooth Amblycerus whiteheacU Kingsolver 

5. Male internal sac with basal pair of sclerite 
spine-shaped 6 

- Male internal sac with basal pair of sclerite Y- 
shaped Amblycerus cistelinus (Gyllenhal) 

6. Male internal sac with unpaired median sclerite 
serrate basally along one side 

. . . Amblycerus sosio Ribeiro-Costa and Kingsolver 

- Male internal sac with unpaired median sclerite 
with serration only on base 

Amblycerus jatayens is (Pic) 

7. Metepisternum with striate area transverse to 
metepisternal sulcus, apparently being a mod- 
ification of metepisternum; apical portion of 
ventral margin of metafemur finely striate, with 
conspicuous tooth; pygidium with median line 



346 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



. . . Amblycerus stridiilator Kingsolver, Romero 

N., and Johnson 

- Metepisternum with striate area running with 
longitudinal axis of metepisternal sulcus, ap- 
parently being a modification of it; apical por- 
tion of ventral margin of metafemur with con- 
spicuous or inconspicuous smooth tooth; py- 
gidium with or without median line 8 

8. Apical portion of ventral margin of metafemur 
with one inconspicuous tooth; posterior end of 
metepisternum straight on striate area .... 

Amblycerus pollens (Sharp) 

- Apical portion of ventral margin of metafemur 
with one conspicuous tooth; posterior end of 
metepisternum curved on striate area 9 

9. Posterior end of metepisternum gently curved 
on striate area; scutellum tridentate; pygidium 
with narrow median line 

Amblycerus eiistrophoides (Schaeffer) 

- Posterior end of metepisternum strongly 
curved on striate area (Fig. 13); scutellum tri- 
dentate on round base (Fig. 12); pygidium oth- 
erwise . . . Amblycerus iscliiodontus. new species 



Acknowledgments 

I thank Dra. Dilma Solange Napp and 
Dra. Catherine N. Duckett for reviewing the 
manuscript. This is Contribution No. 1049 



from Departamento de Zoologia, Universi- 
dade Federal do Parana. 

Literature Cited 

Arrow, G. H. 1904. Sound production in the lamelli- 
corn beetles. Transactions of the Royal Entomo- 
logical Society of London 4: 709-750. 

Dumortier, B. 1963. Morphology of sound emission 
apparatus in Arthropoda, pp. 277-345. In Busnel 
R. G., Acoustic Behavior of Animals. Elsevier 
Publ. Co.. New York. 933 pp. 

Kingsolver, J. M. 1970. A synopsis of the subfamily 
Amblycerinae Bridwell in the West Indies, with 
descriptions of new species (Coleoptera: Bruchi- 
dae). Transactions of the American Entomological 
Society 96: 469-497. 

Kingsolver, J. M., N. J. Romero, and C. D. Johnson. 
1993. Files and scrapers: circumstantial evidence 
for stridulation in three species of Amblycerus, 
one new (Coleoptera: Bruchidae). Pan-Pacific En- 
tomologist 69(2): 122-132. 

Ribeiro-Costa, C. S. 1987. Descri(j6es de oito novas 
especies de Amblycerus Thunberg (Coleoptera: 
Bruchidae). Revista Brasileira de Zoologia 14(3): 
629-648. 

Romero, J., C. D. Johnson, and J. M. Kingsolver. 1996. 
Revision of the genus Amblycerus of the United 
States and Mexico (Coleoptera: Bruchidae: Am- 
blycerinae). United States Department of Agricul- 
ture Technical Bulletin No. 1845, 166 pp. 



PROC. ENTOMOL. SOC. WASH. 
101(2), 1999, pp. 347-358 

A REVIEW OF THE BEACH-FLY GENUS ISOCANACE MATHIS 
(DIPTERA: CANACIDAE) 

Wayne N. Mathis 

Department of Entomology, Smithsonian Institution, Washington, DC 20560-0169, 
U.S.A. (e-mail: mathisw@nmnh.si.edu) 



Abstract. — The species of the beach-fly genus Isocanace Mathis are reviewed, including 
the description of two new species: /. crosbyi (New Zealand. South Island. NN: Cable 
Bay (4r09.6'S, 173°24.9'E) and I.freidbergi (Kenya. Takaungu (50 km N of Mombasa)). 
Descriptions for the genus and two species groups are also revised. 

Key Words: review, Diptera, Canacidae, beach flies, Isocanace, I. crosbyi, I. freidbergi. 
Old World 



Recent field work on New Zealand has 
led to the discovery of the first species of 
the beach-fly family Canacidae from that 
country and has also prompted the research 
that resulted in this review of the genus Is- 
ocanace Mathis. Macrocanace Tonnoir and 
Malloch, with two endemic species from 
New Zealand, had previously been included 
in the Canacidae (Harrison 1959), but is 
now assigned to the family Tethinidae 
(Mathis and Munari 1996). 

The New Zealand species of Canacidae 
is undescribed and is in the genus Isocan- 
ace where it is the second known species 
of the Isocanace albiceps group. The first 
species of that group, /. albiceps (Malloch), 
is from Australia (Mathis 1996). The new 
species is apparently widespread on New 
Zealand, occurring on both North and 
South Islands and undoubtedly on some of 
the associated offshore islands. 

I am also taking this opportunity to de- 
scribe an Afrotropical species of Isocanace 
that Amnon Freidberg and Fini Kaplan col- 
lected some years ago in Kenya. The latter 
species belongs to the Isocanace briani 
group, which previously comprised three 
Afrotropical species: /. australis Mathis 



(Kenya, South Africa), /. briani Mathis (Al- 
dabra, Madagascar), and /. flava (Canzoneri 
and Meneghini; Zaire). The last species, /. 
flava, is unusual among beach flies, being 
one of just a few canacid species that occurs 
in freshwater habitats. As implied by the 
common name for the family, most species 
occur in saline habitats, especially along 
maritime coasts. The new species from 
Kenya is very similar to and is apparently 
the sister species of /. flava, although it is 
found in saline environments along the 
western coast of the Indian Ocean. 

The addition of two new species to Iso- 
canace, one to each of the species groups, 
alters the generic and species-group char- 
acterizations, which are revised. A revised 
key to the species groups and species is also 
provided. 

Isocanace is a relatively recent genus in 
the nomenclatural history of the Canacidae 
(Mathis 1982), and the genus has been 
treated in only two subsequent papers 
(Mathis 1992, 1996). The first paper is a 
world catalog, and the second is a review 
of the Australian beach flies. Isocanace is 
known only from the Afrotropical and Aus- 
tralasian Regions of the Old World (Mathis 



348 



PROCnHDINCiS or THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 



1992). This apparently disjunct distribution 
is probably an artifact due to the poor sam- 
pling of Canacidae from countries between 
Australia and Africa rather than the actual 
distribution of the genus. Nothing is known 
about the immature stages or most other as- 
pects about the biology of the included spe- 
cies except for brief comments on habitats 
where specimens have been collected. 

The descriptive format for the new spe- 
cies follows Mathis and Wirth (1979) and 
Mathis (1982, 1988). More details concern- 
ing the morphology and higher classifica- 
tion of the Canacidae are found in Mathis 
(1982, 1992) and Wirth (1987). I follow 
Crosby et al.'s (1976) geographic codes for 
New Zealand zoogeographic provinces: AK 
= Auckland, NC = North Canterbury, NN 
= Nelson. 

Two venational ratios are used in the de- 
scriptions. Costal vein ratio: The straight 
line distance between the apices of vein 
R2+3 and R4 + ^/distance between the apices 
of veins R, and Ri , <• M vein ratio: The 
straight line distance along vein M between 
crossveins (r-m and dm-cu)/distance apicad 
of dm-cu. 

Specimens are housed in the following 
institutions (acronyms are used in the de- 
scriptive portion of this paper). 

AM Australian Museum, Sydney, Aus- 

tralia 

ANIC The Australian National Insect 
Collection, Division of Entomolo- 
gy, CSIRO, Canberra, Australia 

MNHN Museum National d'Histoire Na- 
turelle, Paris, France 

MR AC Musee Royal de I'Afrique Centra- 
le (Koninklijk Museum voor Mid- 
den-Afrika), Tervuren, Belgium 

NMP Natal Museum, Pietermaritzburg, 
South Africa 

NZAC New Zealand Arthropod Collec- 
tion, Entomology Division, Land- 
care Research, Auckland, New 
Zealand 

TAU Tel Aviv University, Tel Aviv, Is- 
rael 



UQIC University of Queensland Insect 
Collection, Brisbane, Australia 

USNM former United States National Mu- 
seum, collections in the National 
Museum of Natural History, 
Smithsonian Institution, Washing- 
ton, DC, USA 

Genus Isoccmace Mathis 

Isoccmace Mathis 1982: 11. Type species: 
Isocanace briani Mathis, by original des- 
ignation. — Mathis 1992: 5-6 [world cat- 
alog]; 1996: 337-339 [Australian fauna]. 

Canace, in part, of authors: Mathis and 
Wirth 1979: 786. 

Diagnosis. — Resembling Chaetocanace 
Hendel but differing from it and other gen- 
era by the following combination of char- 
acters. 

Head: Mesofrons distinct from para- 
frons, shinier, less microtomentose, with 2— 
3 large, lateral, generally proclinate setae; 
postocellar seta smaller than ocellar seta 
and with more proclinate orientation; 3-4 
pairs of large, lateroclinate, fronto-orbital 
setae; arista plumose, length of branched 
rays varying from approximately subequal 
to nearly twice basal aristal width; dorsally 
genal setae 2-3; dorsally curved genal seta 1. 

Thorax: Dorsocentral setae 4 (1+3); ac- 
rostichal setae evident, in 2 rows, but large, 
prescutellar pair lacking; 2 pairs of scutellar 
setae and frequently some smaller setae in- 
serted dorsally; with only 1 supra-alar seta; 
1-2 notopleural setae, if only 1, anterior 
seta lacking; color of pleural setae variable, 
pale yellow to black; postpronotum bare of 
setulae; katepisternal seta present or absent; 
1 large anepisternal seta; hindtibia lacking 
apical seta anteroventrally; apical section of 
vein M straight. 

Abdomen: Female genital lamellae very 
broad basally, basilateral margins rounded, 
narrowed rather abruptly at level of cleft, 
lamellae very narrow from level of cleft to 
apices, with only 1 large, stout, acute ter- 
minal seta at each apex. Male surstylus 



VOLUME 101, NUMBER 



349 



quite variable, generally slender and with 
apical curvature. 

Distribution. — Old World. Afrotropical 
(Aldabra, Kenya, Madagascar, South Afri- 
ca, and Zaire), Australian (New South 
Wales), New Zealand. 

Discussion. — Mathis and Wirth (1979) 
first noted the possible relationship of some 
species subsequently placed in Isocanace in 
the remarks section of Canace stuckenberi^i 
( = /. hriani). Mathis (1982) concluded that 
this group of species is a monophyletic lin- 
eage that is more closely related to Chae- 
tocanace than to Canace Haliday, sensu 
stricto, and named it as the genus Isocan- 
ace. Study of the type series of Canace fki- 
va Canzoneri and Meneghini revealed that 
this species too is closely related to /. hri- 
ani, thus making a total of four described 
species. The addition of two new species 
brings the number of described species to 
six. 

Key to Species Groui^s and 
Species of Isocanace 

1. Katcpisternal seta lacking; mcsol'rons bare in 
middle; anterior notoplcural seta siiheqiial to 
posterior seta; 2 dorsally genal setae (Austra- 
lasian, the Isoccmace alhiceps group) 2 

- Katcpisternal seta present, although soniclimcs 
pale; mesofrons with scattered selulae on mid- 
dle; anterior notopleural seta distinctly smaller 
than posterior seta or lacking; 3 dorsally genal 
setae (Afrotropical, the /socdiicice hrlcini 
group) 3 

2. Specimens generally more Ian to brown, es- 
pecially Irons, and somewhat on mesonotum 
and dorsum ol abdomen; acroslichal setulae 
sparse and in 2 rows; scutellum conspicuously 
wider than long; surstylus narrow, apex with 
recurved knob (Australia) . . . I. alhiceps (Malloch) 

- Specimens generally gray. Irons appearing blu- 
ish to blackish gray, mesonotum and dorsum 
of abdomen whitish gray to gray, at most par- 
tially tannish medially; acrostichal setulae 
comparatively numerous and in 4 rows, lateral 
row with fewer setulae; scutellum almost as 
long as wide; surstylus broad, apex not re- 
curved or knoblike (New Zealand) 

/. croshyi. new species 

3. Arista with branching rays long, some nearly 
double arislal width at base; anepisternal and 
usually katcpisternal seta pale, mostly yellow- 



ish (Madagascar and Aldabra Islands) .... 

/. hriani Mathis 

Arista with branching rays .shorter, at most 
slightly longer than arislal width at base; ane- 
pisternal and katcpisternal seta black 4 

4. Anterior notopleural seta present, although 
weaker than posterior seta; surstylus in lateral 
view narrow, with subapical posterior swelling 
bearing several pale setae, apical Va curved pos- 
teriorly (Kenya, South Africa) 

/. aiistralis Mathis 

Anterior notopleural seta lacking or very weak 
and pale; surstylus wider in lateral view, with 
subapical anterior swelling, apical Vi narrowed 
considerably and curved anteriorly 5 

5. Lateroclinate fronlo-orbital setae 4. Com- 
pressed sickle-shaped portion of surstylus nar- 
row (Eig. 29) (Zaire) 

/. (lava (Canzoneri and Meneghini) 

- Lateroclinate fronto-orbital setae 3. Com- 
pressed sickle-shaped portion of surstylus 
broad (Fig. 30) (Kenya) 

/. freidhcri^i. new species 



Isocanace albiceps Group 

Diagnosis. — Differing from the Isocan- 
ace hriani group as follows: Head: mesof- 
rons bare in middle; lateroclinate fronto-or- 
bital setae 4; dorsally genal setae 2. Thorax: 
anterior notopleural seta subequal to pos- 
terior seta; katepisternal seta lacking; 
femora concolorous, usually gray to brown- 
ish gray. 

Distribution. — Australasian: Australia, 
New Zealand. 

Isocanace crosbyi Mathis, new species 

(Figs. 1-2) 

Description. — This species is siinilar to /. 
alhiceps but is distinguished from it and 
other congeners by the following combi- 
nation of characters: small to moderately- 
small beach flies, length 1.55-2.15 mm. 

Head: Mesofrons whitish gray to bluish 
gray, subshiny, lacking setulae in middle; 
lateroclinate fronto-orbital setae 4. Antenna 
black with sparse, whitish gray microto- 
mentum; arista bearing short branching 
hairs, these not much longer than basal ar- 
istal width. Gena with 2 large dorsally setae 
and only 2 setulae anterior of ventral dor- 



350 



PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 





Figs. 1-2. External male temiinalia of Isocanace croshyi. 1, Epandrium, cercus, and surstylus, lateral view. 
2, Surstylus, posterior view. 



sally seta, rarely with a posterior setula. 
Maxillary palpus yellow. 

Thorax: Mesonotum brownish gray to 
bluish gray to gray, varying from somewhat 
contrasted to concolorous with grayish 
pleural region; acrostichal setulae in 4 rows, 
lateral row somewhat irregular and with 
fewer setulae; anterior notopleural seta well 
developed, subequal in size and color to 
posterior seta; scutellum almost as long as 
wide. Wing: costal vein ratio 0.15-0.18; M 
vein ratio 0.38-0.45; crossvein r-m slightly 
but consistently basad of middle of discal 
cell. Femora and tibiae gray, concolorous 
with pleural region; tarsi yellow. 

Abdomen: Male terminalia (Figs. 1-2): 



Surstylus in lateral view (Fig. 1) as high as 
epandrium, wider medially, tapered there- 
after to broadly rounded ventral apex; in 
posterior view (Fig. 2) surstylus tapered 
very slightly toward venter until apical Va- 
where medial margin tapered more abrupt- 
ly, making apex narrow and parallel sided. 
Type Material. — The holotype 6 is la- 
beled "NEW ZEALAND. S[OUTH] 
ISL[AND]. NN: Cable Bay (41°09.6'S, 
173°24.9'E), 13 Feb 1998 [,] Wayne N. 
Mathis." The holotype is double mounted 
(minuten in a block of plastic), is in excel- 
lent condition, and is deposited in NZAC. 
Twenty-seven paratypes (24 d, 3 9; 
NZAC, USNM) bear the same label data as 



VOLUME 101, NUMBER 2 



351 



the holotype. Other paratypes are as fol- 
lows: NEW ZEALAND. North Island: AK: 
Auckland Centennial Park (37°0.7'S, 
174°36.3'E), 18 Feb 1998, W. N. Mathis (3 
6, 4 9; USNM); Taramaire Beach 
(37°09.1'S, 175°18.5'E), 8 Feb 1998, W. N. 
Mathis (1 6; USNM). ND: Whananaki 
South (beach; 35°31.1'S, 174°27.2'E), 19 
Feb 1998, W. N. Mathis (19 6, 10 9; 
NZAC, USNM) 

Distribution. — Australasian: New Zea- 
land (AK, NC, NN). 

Etymology. — The species epithet is a 
genitive patronym to recognize the gener- 
ous assistance of Dr. Trevor K. Crosby 
(NZAC) to my research on the shore and 
beach flies of New Zealand. 

Remarks. — There is considerable vari- 
ability in the color of the dorsal surface, 
especially the mesonotum. In the series 
from the type locality, the coloration is 
mostly bluish gray to gray with just faint 
tannish to brownish areas medially. But in 
the long series from Whananaki South there 
is notably more brownish coloration dor- 
sally. The mesofrons, however, remains 
mostly silvery to bluish gray in specimens 
from all localities. 

Isocanace albiceps (Malloch) 
(Figs. 3-12) 

Canace albiceps Malloch 1925: 87 [HT $ 
(AM); Australia. New South Wales: Syd- 
ney]. — Wirth 1951: 262 [review]. 

Isocanace albiceps: Mathis 1982: 18 [ge- 
neric combination]; 1989a: 670 [Austra- 
lasian/Oceanian catalog]; 1992: 6 [world 
catalog]; 1996: 338-339 [Australian fau- 
na]. — Colless and Mc Alpine 1991: 779 
[fig. of head]. 

Diagnosis. — Specimens of /. albiceps are 
similar to those of the Isocanace bri